Apis melliferahoney bee

Ge­o­graphic Range

Apis mel­lif­era is na­tive to Eu­rope, west­ern Asia, and Africa. Human in­tro­duc­tion of Apis mel­lif­era to other con­ti­nents started in the 17th cen­tury, and now they are found all around the world, in­clud­ing east Asia, Aus­tralia and North and South Amer­ica. (Sam­mataro and Avitabile, 1998; Win­ston, et al., 1981)

Habi­tat

Eu­ro­pean hon­ey­bees pre­fer habi­tats that have an abun­dant sup­ply of suit­able flow­er­ing plants, such as mead­ows, open wooded areas, and gar­dens. They can sur­vive in grass­lands, deserts, and wet­lands if there is suf­fi­cient water, food, and shel­ter. They need cav­i­ties (e.g. in hol­low trees) to nest in. (Milne and Milne, 2000; Win­ston, et al., 1981)

Phys­i­cal De­scrip­tion

Gen­er­ally, Apis mel­lif­era are red/brown with black bands and or­ange yel­low rings on ab­domen. They have hair on tho­rax and less hair on ab­domen. They also have a pollen bas­ket on their hind legs. Hon­ey­bee legs are mostly dark brown/black.

There are two castes of fe­males, ster­ile work­ers are smaller (adults 10-15 mm long), fer­tile queens are larger (18-20 mm). Males, called drones, are 15-17 mm long at ma­tu­rity. Though smaller, work­ers have longer wings than drones. Both castes of fe­males have a stinger that is formed from mod­i­fied ovipos­i­tor struc­tures. In work­ers, the sting is barbed, and tears away from the body when used. In both castes, the stinger is sup­plied with venom from glands in the ab­domen. Males have much larger eyes than fe­males, prob­a­bly to help lo­cate fly­ing queens dur­ing mat­ing flights.

There are cur­rently 26 rec­og­nized sub­species of Apis mel­lif­era, with dif­fer­ences based on dif­fer­ences in mor­phol­ogy and mol­e­c­u­lar char­ac­ter­is­tics. The dif­fer­ences among the sub­species is usu­ally dis­cussed in terms of their agri­cul­tural out­put in par­tic­u­lar en­vi­ron­men­tal con­di­tions. Some sub­species have the abil­ity to tol­er­ate warmer or colder cli­mates. Sub­species may also vary in their de­fen­sive be­hav­ior, tongue length, wingspan, and col­oration. Ab­dom­i­nal band­ing pat­terns also dif­fer - some darker and some with more of a mix be­tween darker and lighter band­ing pat­terns.

Hon­ey­bees are par­tially en­dother­mic -- they can warm their bod­ies and the tem­per­a­ture in their hive by work­ing their flight mus­cles. (Clarke, et al., 2002; Milne and Milne, 2000; Pinto, et al., 2004; See­ley, et al., 1982)

  • Sexual Dimorphism
  • female larger
  • sexes shaped differently
  • Range length
    10 to 20 mm
    0.39 to 0.79 in

De­vel­op­ment

Hon­ey­bees build a hive out of wax se­cre­tions from their bod­ies, and queens lay their eggs in cells in the wax. The speed of sub­se­quent de­vel­op­ment of the young is strongly af­fected by tem­per­a­ture, and is fastest at 33-36°C.

Hon­ey­bees are holometabolous in­sects, and have four stages in the life cycle: egg, larva, pupa, and adult.

A. mel­lif­era eggs hatch in 28-144 hours, de­pend­ing on their tem­per­a­ture. The larva that emerges is a small white grub. It stays in its wax cell, grow­ing, and is fed and groomed by adult work­ers. The food that a fe­male larva re­ceives de­ter­mines whether it will be a queen or worker. At 34°C, lar­vae feed and grow for 4-5 days, queens for 6 days, and males for 6-7 days. At the end of that pe­riod their cell is sealed by adult work­ers, and the larva molts, spins a silk co­coon, and trans­forms into the pupa stage. Pupae un­dergo a mas­sive meta­mor­pho­sis that takes about 7-8 days for queens, 12 days for work­ers, and 14-15 days for males. Once their final meta­mor­pho­sis is com­plete, they chew their way out of the cell and begin their adult life. They will not grow or molt after emerg­ing. Adult work­ers will live for 2-4 weeks in the sum­mer, or as long as 11 months if they live through the win­ter. Males only sur­vive for 4-8 weeks, and do not live through the win­ter. Queens live 2-5 years.

. The next stage is the lar­val stage where the larva is fed the royal jelly, pollen/nec­tar, and honey com­bi­na­tion. Next the larva goes into the pupae stage where it caps it­self into its cell to meta­mor­phose into the ma­ture stage.

Queens nor­mally take 16 days to reach ma­tu­rity, the worker bees take 21 days, and the drone takes 24 days to ma­ture. (Ad­jare, 1990; Sam­mataro and Avitabile, 1998)

Re­pro­duc­tion

The great ma­jor­ity of fe­male A. mel­lif­era in a hive are ster­ile work­ers. Only queens mate and lay eggs. Nor­mally there is only a sin­gle re­pro­duc­tive queen in a hive.

Dur­ing pe­ri­ods of suit­ably mild weather in spring and sum­mer, males leave the hive and gather at "drone as­sem­bly areas" near the hive. Vir­gin queens will fly through these areas, at­tract­ing the males with pheromones. The males pur­sue, and at­tempt to mate with the queen in flight. Some­times a "comet" forms, as a clus­ter of males forms around the fe­male, with a string of other males try­ing to catch up. Each male who suc­ceeds in mat­ing drops away, and dies within a few hours or days. Males who do not mate will con­tinue to loi­ter in the as­sem­bly areas until they mate or die try­ing. Queens will mate with up to 10 males in a sin­gle flight.

Queens may mate with males from their own hive, or from other hives in the area. The queen's mat­ing be­hav­ior is cen­tered around find­ing the best place to mate be­fore­hand, by tak­ing di­rec­tional flights for a pe­riod of time, last­ing no more than a cou­ple of days. Af­ter­ward, she leaves the hive and flies to mate with drones in an as­sem­bly area. This nor­mally starts to occur after their first week of birth. The queen does this up to four times. After this con­gre­gate of mat­ing has oc­curred, she never mates again in her life­time. (Ad­jare, 1990; Sam­mataro and Avitabile, 1998; Tarpy and Page Jr., 2000)

Apis mel­lif­era queens are the pri­mary re­pro­duc­ers of the nest and all of the ac­tiv­i­ties of the colony are cen­tered around their re­pro­duc­tive be­hav­iors and their sur­vival. The queen is the only fer­tile fe­male in the colony. She lays eggs nearly con­tin­u­ously through­out the year, some­times paus­ing in late fall in cold cli­mates. A par­tic­u­larly fer­tile queen may lay as many as 1,000 eggs/day, and 200,000 eggs in her life­time. It takes a queen about 16 days to reach adult­hood, and an­other week or more to begin lay­ing eggs. Males take about 24 days to emerge as adults, and begin leav­ing the nest for as­sem­bly areas a few days after that.

Queen hon­ey­bees can con­trol whether or not an egg they lay is fer­til­ized. Un­fer­til­ized eggs de­velop as males and are hap­loid (have only one set of chro­mo­somes). Fer­til­ized eggs are diploid (two sets of chro­mo­somes) and de­velop as work­ers or new queens, de­pend­ing on how they are fed as lar­vae. Queens may in­crease the ratio of male to fe­male eggs they lay if they are dis­eased or in­jured, or in re­sponse to prob­lems in the colony.

Healthy, well-fed hon­ey­bee colonies re­pro­duce by "swarm­ing." The work­ers in the colony begin by pro­duc­ing nu­mer­ous queen lar­vae. Shortly be­fore the new queens emerge, the res­i­dent, egg-lay­ing queen leaves the hive, tak­ing up to half the work­ers with her. This "swarm" forms a tem­po­rary group in a tree nearby, while work­ers scout for a suit­able lo­ca­tion for a new hive. Once they find one, the swarm moves into the space, and be­gins build­ing comb and start­ing the process of food col­lec­tion and re­pro­duc­tion again.

Mean­while at the old hive, the new queens emerge from their cells. If the pop­u­la­tion of work­ers is large enough, and there are few queens emerg­ing, then the first one or two may leave with "af­ter­swarms" of work­ers. After the swarm­ing is com­pleted, any re­main­ing new queens try to sting and kill each other, con­tin­u­ing to fight until all but one is dead. After her com­pe­ti­tion is re­moved, the sur­viv­ing queen be­gins to lay eggs.

Nor­mally the pheromones se­creted by a healthy queen pre­vent work­ers from re­pro­duc­ing, but if a colony re­mains queen­less for long, some work­ers will begin lay­ing eggs. These eggs are un­fer­til­ized, and so de­velop as males. (Ad­jare, 1990; Milne and Milne, 2000; Sam­mataro and Avitabile, 1998; Tarpy and Page Jr., 2000)

  • Breeding interval
    Colonies typically swarm once or twice a year, usually at the beginning of the season that provides the most nectar.
  • Breeding season
    Late spring until the winter months
  • Range eggs per season
    60,000 to 80,000
  • Average gestation period
    3 days
  • Range age at sexual or reproductive maturity (female)
    15 to 17 days
  • Average age at sexual or reproductive maturity (male)
    24 days

As in most eu­so­cial in­sects, the off­spring of fer­tile fe­males (queens) are cared for other mem­bers of the colony. In hon­ey­bees, the care­tak­ers are ster­ile fe­males, daugh­ters of the queen, called work­ers.

Work­ers build and main­tain the comb where young bees are raised, gather food (nec­tar and pollen) feed and tend lar­vae, and de­fend the hive and its young from preda­tors and par­a­sites.

Young queens in­herit their hive from their moth­ers. Often sev­eral new queens emerge after the old queen leaves with a swarm to found a new colony. The new queens fight for con­trol of the hive, and only one sur­vives the con­flict. (Ad­jare, 1990; Sam­mataro and Avitabile, 1998)

  • Parental Investment
  • pre-fertilization
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female
  • inherits maternal/paternal territory
  • maternal position in the dominance hierarchy affects status of young

Lifes­pan/Longevity

Apis mel­lif­era queens usu­ally live 2 to 3 years, but some have been known to last for 5 years. Work­ers typ­i­cally only live for a few weeks, some­times a few months if their hive be­comes dor­mant in win­ter. Males live for 4-8 weeks at the most. (Tarpy and Page Jr., 2000)

  • Typical lifespan
    Status: wild
    2 to 3 years

Be­hav­ior

Eu­ro­pean hon­ey­bees are eu­so­cial in­sects. They live in colonies that con­tain one re­pro­duc­tive fe­male (the queen) and her off­spring. Ster­ile fe­male off­spring of the queen (the work­ers) per­form all the work of the colony and are by far the most nu­mer­ous caste in the hive. Males and queens spend all their ef­fort on re­pro­duc­tion, see the Re­pro­duc­tion sec­tions for in­for­ma­tion on mat­ing be­hav­ior and egg pro­duc­tion.

A. mel­lif­era work­ers show what is called "age poly­ethism." Their be­hav­iors change as they get older. Newly emerged work­ers clean cells, prepar­ing them for a new egg or for food stor­age. After a few days they shift to other hive main­te­nance work, re­mov­ing waste and de­bris, fan­ning to main­tain air cir­cu­la­tion and tem­per­a­ture, pro­cess­ing nec­tar brought by for­agers, and feed­ing the queen and lar­vae from glands in their head and body. In their sec­ond week of adult life work­ers' wax glands be­come ac­tive and they help build and re­pair the comb, while con­tin­u­ing to tend the queen and feed work­ers. Apis mel­lif­era work­ers build a "comb", a sheet of hexag­o­nal cells made of waxes they se­crete. Each cell can house one lar­val bee, and cells are also used as pro­tected stor­age space for honey (processed nec­tar) and pollen.

Be­tween 12 and 25 days, work­ers take a turn guard­ing the hive, in­spect­ing any bees that try to enter the hive - dri­ving off strangers and at­tack­ing any other crea­tures that try to enter. After about three weeks, the work­ers food and wax glands at­ro­phy, and they shift to for­ag­ing duty.

For­ag­ing only oc­curs dur­ing day­light, but bees are ac­tive in the hive con­tin­u­ously.

In tem­per­ate cli­mates, colonies store honey and pollen to feed on dur­ing the win­ter. Dur­ing cold tem­per­a­tures the work­ers and queen form a tight ball or clus­ter, work­ing their flight mus­cles to gen­er­ate heat and keep them­selves warm. In warmer trop­i­cal re­gions, hon­ey­bees main­tain smaller stores of food.

If a colony's nest con­di­tions be­come too poor, the en­tire colony may move to a new site. This is par­tic­u­larly com­mon in trop­i­cal hon­ey­bees, that move in re­sponse to sea­sonal drought. Bee­keep­ers call this "ab­scond­ing", and work to pre­vent it in do­mes­ti­cated colonies.

Swarm­ing is a be­hav­ior in a nest where a new queen is born that takes the place of the older one in that hive. The de­part­ing queen nor­mally takes some of the work­ers with her. Swarm­ing bees send out worker scouts to look for a suit­able home to take the place of the one they left. The swarm of bees is just tem­po­rary. They nor­mally swarm over a twig or branch of a tree or any­where that can be used tem­porar­ily as an in­ter­me­di­ate nest. (Ad­jare, 1990; Sam­mataro and Avitabile, 1998)

Home Range

Hon­ey­bees for­age as close to the hive as pos­si­ble, usu­ally within a 3 kilo­me­ter ra­dius around the hive (i.e. an area of about about 2800 hectares). If nec­es­sary, they can fly as far as 8-13 km to reach food or water. (Per­ci­val, 1947; Sam­mataro and Avitabile, 1998)

Com­mu­ni­ca­tion and Per­cep­tion

Apis mel­lif­era com­mu­ni­ca­tion is based on chem­i­cal sig­nals, and most of their com­mu­ni­ca­tion and per­cep­tion be­hav­iors are cen­tered around scent and taste. The mem­bers of the hive colony are bound chem­i­cally to each other. Each hive has a unique chem­i­cal sig­na­ture that hive­mates use to rec­og­nize each other and de­tect bees from other colonies.

Within the hive, bees are in con­stant chem­i­cal com­mu­ni­ca­tion with each other. Work­ers feed and groom each other, as well as lar­vae, drones, and the queen. In the process they pass on pheromones, chem­i­cal sig­nals that in­di­cate in­for­ma­tion about the health of the queen and the state of the colony.

Chem­i­cals not only help with de­tect­ing the right sig­na­ture of hives but also with for­ag­ing. Hon­ey­bees use scent to lo­cate flow­ers from a dis­tance. When a suc­cess­ful for­ager re­turns to the hive, it passes the scent of the flow­ers to its nest mates, to help them find the same patch of flow­ers.

Bees also use chem­i­cals to sig­nal out­side the hive. When a worker stings some­thing, her stinger re­leases an alarm pheromone that causes other bees to be­come ag­i­tated, and helps them lo­cate the enemy.

Thought it's al­ways dark in the hive, vi­sion is im­por­tant to hon­ey­bees out­side. They can see other an­i­mals, and rec­og­nize flow­ers. The eyes of Apis species can de­tect ul­tra­vi­o­let light wave­lengths that are be­yond the vis­i­ble spec­trum. This al­lows them to lo­cate the sun on cloudy days, and see mark­ings on flow­ers that are only vis­i­ble in ul­tra­vi­o­let light. One por­tion of hon­ey­bee's eyes is sen­si­tive to po­lar­ized light, and they use this to nav­i­gate.

Work­ers and queens can hear vi­bra­tions. New queens call to each other and work­ers when they first emerge. Work­ers hear the vi­bra­tions of the wag­gle dances made by re­turn­ing for­agers.

Apis species have a par­tic­u­larly no­table form of com­mu­ni­ca­tion called "danc­ing." For­agers that have lo­cated an abun­dant sup­ply of food do a dance to com­mu­ni­cate the lo­ca­tion of the patch to other for­agers. A "round dance" in­di­cates food within about 300 me­ters of the hive, and only com­mu­ni­cates the pres­ence of the flow­ers, not the di­rec­tion, though work­ers will also get the scent from the food the for­ager has brought back. The more com­pli­cated "wag­gle dance" in­di­cates the di­rec­tion and dis­tance of food fur­ther away, using the lo­ca­tion of the sun and the bee's mem­ory of the dis­tance it flew to re­turn to the hive. Sym­bolic com­mu­ni­ca­tion is quite un­usual among in­ver­te­brates, and these hon­ey­bee "dances" have been in­ten­sively stud­ied. (Breed, et al., 1985; Milne and Milne, 2000; Rein­hard, et al., 2004; Roat and Landim, 2008; Sam­mataro and Avitabile, 1998; San­doz, et al., 2002; Sher­man and Viss­cher, 2002)

Food Habits

Apis mel­lif­era feed on pollen and nec­tar col­lected from bloom­ing flow­ers. They also eat honey (stored, con­cen­trated nec­tar) and se­cre­tions pro­duced by other mem­bers of their colony.

Work­ers for­age for food (nec­tar and pollen) for the en­tire colony. They use their tongues to suck up nec­tar, and store it in the an­te­rior sec­tion of the di­ges­tive tract, called the crop. They col­lect pollen by groom­ing it off the bod­ies and onto spe­cial struc­tures on their hind legs called pollen bas­kets.

Re­turn­ing for­agers trans­fer the nec­tar they have col­lected to younger worker bees that in turn feed other mem­bers of the hive, or process it into honey for long-term stor­age. They add en­zymes to the honey, and store it in open cells where the water can evap­o­rate, con­cen­trat­ing the sug­ars.

Young work­ers eat pollen and nec­tar, and se­crete food ma­te­ri­als, called “royal jelly” and “worker jelly”, from glands in their heads. This ma­te­r­ial is fed to young lar­vae, and the amount and type they get de­ter­mines if they will be queens or work­ers.

Hon­ey­bees for­age dur­ing day­light hours, but are equally ac­tive on cloudy or sunny days. They will not fly in heavy rain or high winds, or if the tem­per­a­ture is too ex­treme (work­ers can't fly when they get below 10°C). Dur­ing the warm, calm weather the hon­ey­bees col­lect the most pollen even if it is cloudy. If the light in­ten­sity changes rapidly, they im­me­di­ately stop work­ing and re­turn to the hive. If it lightly rains, pollen col­lec­tion stops, be­cause mois­ture in­hibits the bee’s abil­ity to col­lect it. How­ever, nec­tar col­lec­tion is not in­hib­ited by light rain. Wind also af­fects the rate of pollen col­lec­tion.

Hon­ey­bee work­ers are op­por­tunis­tic. They will steal from other hives if they can. Hive-rob­bing can be dan­ger­ous, but a weak­ened or dam­aged hive may be raided by work­ers from other hives, es­pe­cially when nec­tar flows in flow­ers are not abun­dant. Hon­ey­bees will also col­lect “hon­ey­dew,” the sweet fluid ex­creted by sap-feed­ing in­sects like aphids. (Ad­jare, 1990; Gon­za­lez, et al., 1995; Per­ci­val, 1947; Sam­mataro and Avitabile, 1998)

  • Plant Foods
  • nectar
  • pollen
  • sap or other plant fluids

Pre­da­tion

Hon­ey­bees have many adap­ta­tions for de­fense: Adults have or­ange and black strip­ing that acts as warn­ing col­oration. Preda­tors can learn to as­so­ci­ate that pat­tern with a painful sting, and avoid them. Hon­ey­bees pre­fer to build their hives in pro­tected cav­i­ties (small caves or tree hol­lows). They seal small open­ings with a mix of wax and resins called propo­lis, leav­ing only one small open­ing. Worker bees guard the en­trance of the hive. They are able to rec­og­nize mem­bers of their colony by scent, and will at­tack any non-mem­bers that try to enter the hive. Work­ers and queens have a ven­omous sting at the end of the ab­domen. Un­like queens, and un­usual among sting­ing in­sects, the stings of Apis work­ers are heav­ily barbed and the sting and venom glands tear out of the ab­domen, re­main­ing em­bed­ded in the tar­get. This causes the death of the worker, but may also cause a more painful sting, and dis­cour­age the preda­tor from at­tack­ing other bees or the hive. A sting­ing worker re­leases an alarm pheromone which causes other work­ers to be­come ag­i­tated and more likely to sting, and sig­nals the lo­ca­tion of the first sting.

Hon­ey­bees are sub­ject to many types of preda­tors, some at­tack­ing the bees them­selves, oth­ers con­sum­ing the wax and stored food in the hive. Some preda­tors are spe­cial­ists on bees, in­clud­ing hon­ey­bees.

Im­por­tant in­ver­te­brate en­e­mies of adult bees in­clude crab spi­ders and orb-weaver spi­ders, wasps in the genus Phil­an­thus (called “bee­wolves”), and many species of so­cial wasps in the fam­ily Vesp­i­dae. Vespid wasp colonies are known to at­tack hon­ey­bee colonies en masse, and can wipe out a hive in one at­tack. Many ver­te­brate in­sec­ti­vores also eat adult hon­ey­bees. Toads (Bufo) that can reach the en­trance of hive will sit and eat many work­ers, as will opos­sums (Didel­phis). Birds are an im­por­tant threat – the Merop­i­dae (bee-eaters) in par­tic­u­lar in Africa and south­ern Eu­rope, but also fly­catch­ers around the world (Tyrranidae and Mus­c­i­cap­i­dae). Apis mel­lif­era in Africa are also sub­ject to at­tack by hon­eyguides. These birds eat hive comb, con­sum­ing bees, wax, and stored honey. At least one species, the greater hon­eyguide (In­di­ca­tor in­di­ca­tor) will guide mam­mal hive preda­tors to hives, and then feed on the hive after the mam­mal has opened it up.

The main ver­te­brate preda­tors of hives are mam­mals. Bears fre­quently at­tack the nests of so­cial bees and wasps, as do many mustelids such as the tayra in the Neotrop­ics and es­pe­cially the honey bad­ger of Africa and south­ern and west­ern Asia. In the West­ern Hemi­sphere skunks, ar­madil­los and anteaters also raid hives, as do pan­golins (Manis) in Africa. Large pri­mates, in­clud­ing ba­boons, chim­panzees (<<g.​Pan>>) and go­ril­las are re­ported to at­tack hives too. Smaller mam­mals such as mice (Mus) and rats (Rat­tus) will bur­row into hives as well.

Some in­sects are preda­tors in hives as well, in­clud­ing wax moth lar­vae (Gal­le­ria mel­lonella, Achroia grisella), and hive bee­tles (Hy­lostoma, Aethina), and some species of ants. In their na­tive re­gions these tend not to be im­por­tant en­e­mies, but where hon­ey­bees have not co-evolved with these in­sects and have no de­fense, they can do great harm to hives.

See Ecosys­tem Roles sec­tion for in­for­ma­tion on hon­ey­bee par­a­sites and pathogens. (Ad­jare, 1990; Roubik, 1989; Sam­mataro and Avitabile, 1998)

Ecosys­tem Roles

Hon­ey­bees are very im­por­tant pol­li­na­tors, and are the pri­mary pol­li­na­tor for many plants. With­out hon­ey­bees, these plants have greatly re­duced fer­til­ity. In North Amer­ica and Aus­tralia, where there are no na­tive bee species with large colonies, hon­ey­bees can have es­pe­cially strong ef­fects on na­tive flow­ers, and on other pol­li­na­tors such as soli­tary bee species. Hon­ey­bees abil­ity to re­cruit fel­low work­ers by “danc­ing” al­lows them to be more ef­fi­cient than other pol­li­na­tors at ex­ploit­ing patches of flow­ers. This can cre­ate strong im­pacts on their com­peti­tors, es­pe­cially soli­tary bees.

Like all so­cial in­sects, hon­ey­bees are hosts to a va­ri­ety of par­a­sites, com­men­sal or­gan­isms, and path­o­genic mi­crobes. Some of these can be se­ri­ous prob­lems for api­cul­ture, and have been stud­ied in­ten­sively. At least 18 types of viruses have been found to cause dis­ease in bees, in­clud­ing Sacbrood dis­ease. Sev­eral of them (but not sacbrood virus) are as­so­ci­ated with par­a­sitic mites. Bac­te­ria in­fect bees, no­tably Bacil­lus lar­vae, agent of Amer­i­can Foul­brood dis­ease, and Melis­so­coc­cus plu­ton, agent of Eu­ro­pean Foul­brood. Fungi grow in bee hives, and As­cosphaera apis can cause Chalk­brood dis­ease. One of the most com­mon dis­eases in do­mes­ti­cated hives is Nosema dis­ease, caused by a pro­to­zoan, Nosema apis. An amoeba, Mal­phig­amoeba mel­li­fi­cae, also causes dis­ease in hon­ey­bees.

In re­cent decades, two mite species have spread through do­mes­ti­cated and feral hon­ey­bee pop­u­la­tions around the world. Acara­pis woodi is a small mite species that lives in the tra­cheae of adult bees and feeds on bee he­molymph. It was first dis­cov­ered in Eu­rope, but its ori­gin is un­known. In­fes­ta­tions of these mites weaken bees, and in cold cli­mates, whole colonies may fail when the bees are con­fined in the hive dur­ing the win­ter. A much worse threat is Var­roa de­struc­tor. This might evolved on an Asian hon­ey­bee, Apis cer­ana, but switched on to Apis mel­lif­era colonies that were set up in east Asia. It has since spread all around the world, ex­cept Aus­tralia. Ju­ve­nile mites feed on bee lar­vae and pupae, and adult fe­male mites feed and dis­perse on adult work­ers. This mite is known to spread sev­eral viruses as well. In­fes­ta­tions of V. de­struc­tor often wipe out colonies. Nearly all the feral, un­tended hon­ey­bee colonies in North Amer­i­can are be­lieved to have been wiped out by mite in­fes­ta­tions, along with a large pro­por­tion of do­mes­ti­cated colonies. Other mite species are known from hon­ey­bee colonies, but they are not con­sid­ered harm­ful.

An­other com­men­sal or par­a­sitic species is Braula coeca, the bee louse. De­spite the com­mon name, this is ac­tu­ally a wing­less fly, that ap­par­ently feeds by in­ter­cept­ing food being trans­ferred from one bee to an­other.

Bee­tles in the gen­era Hy­lostoma and Aethina are found in African hon­ey­bee nests, where they seem to do lit­tle harm. How­ever, the "small hive bee­tle", Aethina tu­mida, has be­come a sig­nif­i­cant prob­lem in Eu­ro­pean and North Amer­i­can hives. The lar­vae eat all the con­tents of comb: honey, pollen, and bee eggs and lar­vae. (Ad­jare, 1990; Roubik, 1989; Sam­mataro and Avitabile, 1998)

Com­men­sal/Par­a­sitic Species

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Hon­ey­bees pol­li­nate bil­lions of US dol­lars worth of com­mer­cial agri­cul­tural crops around the world every year. They are im­por­tant pol­li­na­tors for eco­nom­i­cally im­por­tant wild plant pop­u­la­tions as well.

Hon­ey­bee hives pro­vide honey and wax, and pollen, propo­lis, and royal jelly that are sold for med­i­cines and cos­met­ics.

Hon­ey­bees are im­por­tant study or­gan­isms for re­search in the con­nec­tions be­tween ner­vous sys­tem struc­ture and be­hav­ior.

Some re­search sug­gests hon­ey­bee venom may have med­ically use­ful ap­pli­ca­tions in the treat­ment of auto-im­mune dis­ease or in­flam­ma­tion. (Ad­jare, 1990; Kang, et al., 2002; Sam­mataro and Avitabile, 1998)

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Hon­ey­bee work­ers will sting hu­mans and do­mes­ti­cated an­i­mals in de­fense of them­selves or their hive. A sin­gle sting is painful but not dan­ger­ous un­less the tar­get is al­ler­gic to the venom, in which case it can be life threat­en­ing. Oth­er­wise, it takes about 20 stings per kilo­gram of body weight to be life threat­en­ing.

Each sub­species of Apis mel­lif­era has dif­fer­ent be­hav­ioral pat­terns in re­gards to in­trud­ers near or around the hive. The African sub­species are par­tic­u­larly ag­gres­sive. One of them, Apis mel­lif­era scutel­lata, was ac­ci­den­tally re­leased in South Amer­ica, and has spread north to the south­ern United States. This is the "killer bee." It is no­table for hav­ing a much higher ag­gres­sive re­sponse to dis­tur­bance -- more work­ers at­tack than in other sub­species, and they pur­sue tar­gets much longer than Eu­ro­pean bees do. The spread of these bees made bee­keep­ing much more ex­pen­sive and com­pli­cated, and the ag­gres­sive bees caused many deaths. (Ad­jare, 1990; Sam­mataro and Avitabile, 1998)

  • Negative Impacts
  • injures humans

Con­ser­va­tion Sta­tus

While the species as a whole is still very nu­mer­ous, there is con­cern in Eu­rope that wide­spread com­mer­cial­iza­tion of bee­keep­ing is en­dan­ger­ing lo­cally-adapted pop­u­la­tions and sub­species. This, com­bined with higher mor­tal­ity of colonies due to Var­roa mite and tra­cheal mite in­fes­ta­tions, and the re­cent phe­nom­e­non of Colony Col­lapse Dis­or­der in North Amer­ica, has cause sig­nif­i­cant con­cern for the health of the pop­u­la­tion. Colony Col­lapse Dis­or­der (CCD) is a con­di­tion of com­mer­cial bee­hives, where there are sud­den mas­sive waves of mor­tal­ity among the work­ers. Bee­keep­ers dis­cover their hives sim­ply empty of work­ers, with so few sur­viv­ing that they can­not tend the queen and brood. This con­di­tion has oc­curred mainly in North Amer­ica, and mainly in large com­mer­cial api­aries. No sin­gle cause has been iden­ti­fied yet. (Ad­jare, 1990; Sam­mataro and Avitabile, 1998)

Con­trib­u­tors

George Ham­mond (au­thor, ed­i­tor), An­i­mal Di­ver­sity Web, Madi­son Blanken­ship (au­thor), Rad­ford Uni­ver­sity, Karen Pow­ers (ed­i­tor, in­struc­tor), Rad­ford Uni­ver­sity.

Glossary

Australian

Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.

World Map

Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

World Map

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

aposematic

having coloration that serves a protective function for the animal, usually used to refer to animals with colors that warn predators of their toxicity. For example: animals with bright red or yellow coloration are often toxic or distasteful.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chaparral

Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.

chemical

uses smells or other chemicals to communicate

colonial

used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.

cosmopolitan

having a worldwide distribution. Found on all continents (except maybe Antarctica) and in all biogeographic provinces; or in all the major oceans (Atlantic, Indian, and Pacific.

desert or dunes

in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.

diurnal
  1. active during the day, 2. lasting for one day.
drug

a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

eusocial

the condition in which individuals in a group display each of the following three traits: cooperative care of young; some individuals in the group give up reproduction and specialize in care of young; overlap of at least two generations of life stages capable of contributing to colony labor

fertilization

union of egg and spermatozoan

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

herbivore

An animal that eats mainly plants or parts of plants.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

hibernation

the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.

induced ovulation

ovulation is stimulated by the act of copulation (does not occur spontaneously)

internal fertilization

fertilization takes place within the female's body

introduced

referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

keystone species

a species whose presence or absence strongly affects populations of other species in that area such that the extirpation of the keystone species in an area will result in the ultimate extirpation of many more species in that area (Example: sea otter).

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nectarivore

an animal that mainly eats nectar from flowers

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

polarized light

light waves that are oriented in particular direction. For example, light reflected off of water has waves vibrating horizontally. Some animals, such as bees, can detect which way light is polarized and use that information. People cannot, unless they use special equipment.

polyandrous

Referring to a mating system in which a female mates with several males during one breeding season (compare polygynous).

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

seasonal breeding

breeding is confined to a particular season

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

sperm-storing

mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.

stores or caches food

places a food item in a special place to be eaten later. Also called "hoarding"

suburban

living in residential areas on the outskirts of large cities or towns.

swamp

a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

urban

living in cities and large towns, landscapes dominated by human structures and activity.

venomous

an animal which has an organ capable of injecting a poisonous substance into a wound (for example, scorpions, jellyfish, and rattlesnakes).

vibrations

movements of a hard surface that are produced by animals as signals to others

visual

uses sight to communicate

Ref­er­ences

Abreu, R., S. Moraes, O. Malaspina. 2000. His­to­log­i­cal as­pects and pro­tein con­tent of Apis mel­lif­era L. worker venom glands: the ef­fect of elec­tri­cal shocks in sum­mer and win­ter. Jour­nal of Ven­omous An­i­mals and Tox­ins, 6/1: 87-98.

Ad­jare, S. 1990. Bee­keep­ing in Africa. Rome, Italy: Food and Agri­cul­ture Or­gan­i­sa­tion of the United Na­tions. Ac­cessed No­vem­ber 06, 2008 at http://​www.​fao.​org/​docrep/​t0104e/​T0104E00.​htm.

Amdam, G., K. Nilsen, K. Nor­berg, M. Fon­drk, K. Hart­felder. 2007. Vari­a­tion in en­docrine sig­nal­ing un­der­lies vari­a­tion in so­cial life his­tory. The Amer­i­can Nat­u­ral­ist, 170/1: 37-46.

Breed, M., L. But­ler, T. Stiller. 1985. Kin dis­crim­i­na­tion by worker honey bees in ge­net­i­cally mixed groups. Pro­ceed­ings of the Na­tional Acad­emy of Sci­ences of the United States of Amer­ica, 82/9: 3058-3061.

Clarke, K., T. Rinderer, P. Franck, Q. Javier, B. Ol­droyd. 2002. The african­iza­tion of hon­ey­bees (Apis mel­lif­era L.) of the Yu­catan: a study of a mas­sive hy­bridiza­tion event across time. Evo­lu­tion, 56/7: 1462-1474.

Gon­za­lez, A., C. Rowe, P. Weeks, D. Whit­tle, F. Gilbert, C. Barnard. 1995. Flower choice by honey bees (Apis mel­lif­era L.): sex-phase of flow­ers and pref­er­ences among nec­tar and pollen for­agers. Oe­colo­gia, 101/2: 258-264.

Hem­mer, W., M. Focke, D. Ko­larich, I. Wil­son, F. Alt­mann, S. Wöhrl, M. Götz, R. Jarisch. 2001. An­ti­body bind­ing to venom car­bo­hy­drates is a fre­quent cause for dou­ble pos­i­tiv­ity to hon­ey­bee and yel­low jacket venom in pa­tients with sting­ing-in­sect al­lergy. Jour­nal of Al­lergy and Clin­i­cal Im­munol­ogy, 108/6: 1045-1052.

Kang, S., C. Pak, H. Choi. 2002. The ef­fect of whole bee venom on arthri­tis. The Amer­i­can Jour­nal of Chi­nese Med­i­cine, 30/1: 73-80.

LIPPS, B. 2002. Sub-lethal in­jec­tion of hon­ey­bee venom de­creased the lev­els of en­doge­nously pre­sent sub­stance in or­gans of mice. Jour­nal of Ven­omous An­i­mals and Tox­ins, 8/2: 255-268.

Milne, M., L. Milne. 2000. Na­tional Audubon So­ci­ety: Field Guide To In­sects and Spi­ders. New York, Canada: Al­fred A. Knopf, Inc..

Morse, R. 1978. Honey bee pests, preda­tors, and dis­eases. Ithaca, New York, USA: Cor­nell Uni­ver­sity Press.

Per­ci­val, M. 1947. Pollen col­lec­tion by Apis mel­lif­era . New Phy­tol­o­gist, 46/1: 142-173.

Pinto, A., W. Ru­bink, R. Coul­son, J. Pat­ton, S. John­ston. 2004. Tem­po­ral pat­tern of african­iza­tion in a feral hon­ey­bee pop­u­la­tion from texas in­ferred from mi­to­chon­dr­ial DNA. Evo­lu­tion, 58/5: 1047-1055.

Rein­hard, J., M. Srini­vasan, S. Zhang. 2004. Scent-trig­gered nav­i­ga­tion in hon­ey­bees. Na­ture, 427: 411.

Roat, T., C. Landim. 2008. Tem­po­ral and mor­pho­log­i­cal dif­fer­ences in post-em­bry­onic dif­fer­en­ti­a­tion of the mush­room bod­ies in the brain of work­ers, queens and drones of Apis mel­lif­era (Hy­menoptera: Ap­i­dae). Mi­cron, 39: 1171-1178.

Roubik, D. 1989. Ecol­ogy and nat­ural his­tory of trop­i­cal bees. New York City, New York, USA: Cam­bridge Uni­ver­sity Press.

Sam­mataro, D., A. Avitabile. 1998. The Bee­keeper's Hand­book, 3rd edi­tion. Ithaca, New York, USA: Com­stock Pub­lish­ing As­so­ci­ates.

San­doz, C., M. Ham­mer, R. Men­zel. 2002. Side speci­ficity of ol­fac­tory learn­ing in the hon­ey­bee: US input side. Learn­ing and Mem­ory, 9: 337-348.

See­ley, T., R. See­ley, P. Akratanakul. 1982. Colony de­fense strate­gies of the hon­ey­bees in Thai­land. Eco­log­i­cal Mono­graphs, 52/1: 43-63.

Shemesh, Y., M. Cohen, G. Bloch. 2007. Nat­ural plas­tic­ity in cir­ca­dian rhythms is me­di­ated by re­or­ga­ni­za­tion in mol­e­c­u­lar clock­work in hon­ey­bees. The FASEB Jour­nal, 21: 2304-2311.

Sher­man, G., K. Viss­cher. 2002. Hon­ey­bee colonies achieve fit­ness through danc­ing. Na­ture, 419: 920-922.

South­wick, E., G. Held­maier. 1987. Tem­per­a­ture con­trol in hon­ey­bee colonies. Bio­Science, 37/6: 395-399.

Tarpy, D., R. Page Jr.. 2000. No be­hav­ior con­trol over mat­ing fre­quency in queen hon­ey­bees (Apis mel­lif­era L.): im­pli­ca­tions for the evo­lu­tion of ex­treme polyandry. The Amer­i­can Nat­u­ral­ist, 155/6: 820-827.

Win­ston, M., J. Drop­kin, O. Tay­lor. 1981. De­mog­ra­phy and life his­tory char­ac­ter­is­tics of two honey bee races (Apis mel­lif­era). Oe­colo­gia, 48: 407-413.