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Drone production

Drones usually are not present in the colony during the whole year. In moderate climate they can be found in most colonies during spring and summer The honeybee queen and her attendants,
Anim. Behav.
, Volume 8, p.201-208, (1960)
[1]. Only in queenless colonies drones can be present during autumn and winter. Colonies with larger number of workers start drone production earlier The production of drone comb by honeybee colonies,
J. Apic. Res.
, Volume 6, p.29-36, (1967)
[2]. Initiation of drone rearing occurs about three weeks before swarming but this time varies greatly between colonies Drone Brood in Honey Bee Colonies,
Journal of Economic Entomology
, Volume 51, p.46-48, (1958)
[3]. Maximum number of drones occurs during swarming season The effect of a plentiful supply of drone comb on colonies of honeybees,
J. Apic. Res.
, Volume 4, p.109-119, (1965)
[4]Factors determining the rearing and rejection of drones by the honeybee colony,
Anim. Behav.
, Volume 23, p.650-675, (1975)
[5]Protandrous reproduction in honey bees,
Envir. Entomol.
, Volume 10, p.359-361, (1981)
[6]Effects of reproductive timing and colony size on the survival, offspring colony size and drone production in the honey bee (Apis mellifera),
Ecological Entomology
, Volume 12, Number 2, p.187–195, (1987)
[7]Drone production and drone comb utilization in colonies of the African honey bee, Apis mellifera scutellata Lepeletier, in Africa,
Apidologie
, Volume 25, p.547–547, (1994)
[8]. At this time of year ratio of drone to worker brood is particularly high in swarming colonies Protandrous reproduction in honey bees,
Envir. Entomol.
, Volume 10, p.359-361, (1981)
[6]. In spring workers remove honey from drone cells in order to make space for drone brood Adaptive tuning of an extended phenotype: honeybees seasonally shift their honey storage to optimize male production,
Animal Behaviour
, Volume 103, p.29–33, (2015)
[9]. Peak of drone production can occur at different time of year in different subspecies Partial seasonal isolation of African and European-derived Apis mellifera (Hymenoptera: Apidae) drones at congregation areas from subtropical Mexico,
Annals of the Entomological Society of America
, Volume 94, Number 4, p.540–544, (2001)
[10]. There is no correlation between time of production of first drones and time of swarming Observations on colonies of honey-bees subjected to treatments designed to induce swarming,
Proc. Roy. Entomol. Soc. London (A)
, Volume 32, p.185-192, (1957)
[11]The production of queen cups and queen cells in relation to the general development of honeybee colonies and its connection with swarming and supersedure,
J. Apic. Res.
, Volume 4, p.121-141, (1965)
[12]. Drone brood area changes during a season in similar way to worker brood area The production of queen cups and queen cells in relation to the general development of honeybee colonies and its connection with swarming and supersedure,
J. Apic. Res.
, Volume 4, p.121-141, (1965)
[12]. Later during season production of drones is smaller and in queenright colonies they are not produced later than late summer Reproduction and growth of temperate-evolved honey bee colonies (Apis mellifera L.),
, Burnaby, (1985)
[13]. At this time in temperate climate workers stop feeding drones, deny them access to honey stores and drag them from the nest (see: Drone eviction) Factors determining the rearing and rejection of drones by the honeybee colony,
Anim. Behav.
, Volume 23, p.650-675, (1975)
[5]. Under starvation conditions production of drones is smaller Nutritional factors affecting the egg sex ratio adjustment by a honeybee queen,
Insect Soc
, Volume 48, p.355-359, (2001)
[14] and they are removed from colony faster Fall death rates of drone honey bees,
J. Econ. Entomol.
, Volume 60, p.1198-1202, (1967)
[15].

In natural conditions number of drones produced by one non-swarming colony during one year is about 20 thousand The effect of a plentiful supply of drone comb on colonies of honeybees,
J. Apic. Res.
, Volume 4, p.109-119, (1965)
[4]Protandrous reproduction in honey bees,
Envir. Entomol.
, Volume 10, p.359-361, (1981)
[6] (the exact estimate from one study was 22560 ± 8280 Protandrous reproduction in honey bees,
Envir. Entomol.
, Volume 10, p.359-361, (1981)
[6]). In managed colonies the number of drones produced during one season is smaller (between few hundred and few thousand) because of small number of drone cells which strongly affect production of drones Drone production in honey-bee colonies (Apis mellifera L.),
Nature
, Volume 199, p.789-790, (1963)
[16]The production of queen cups and queen cells in relation to the general development of honeybee colonies and its connection with swarming and supersedure,
J. Apic. Res.
, Volume 4, p.121-141, (1965)
[12]. If large amount of drone comb is provided up to 45 thousand of drones can be produced by one colony. Queenless colony can produce more than six thousand drones Protandrous reproduction in honey bees,
Envir. Entomol.
, Volume 10, p.359-361, (1981)
[6]Reproduction by worker honeybees (Apis mellifera L),
Behav Ecol Sociobiol
, Volume 23, p.117-126, (1988)
[17]. Queenless colonies with large number of patrilines produce fewer drones Genotypic diversity in queenless honey bee colonies reduces fitness,
Journal of Apicultural Research
, Volume 51, Number 4, p.336–341, (2012)
[18]. Colonies with bigger honey reserves produce more drones Reproductive decisions by honey bee colonies: tuning investment in male production in relation to success in energy acquisition,
Insectes Sociaux
, Volume 50, Number 2, p.134-138, (2003)
[19]. Small colonies in poor environmental conditions can produce no drones during whole season The production of queen cups and queen cells in relation to the general development of honeybee colonies and its connection with swarming and supersedure,
J. Apic. Res.
, Volume 4, p.121-141, (1965)
[12]. Drone comb is build only in colonies with more than 4000 workers A critical number of workers in a honeybee colony triggers investment in reproduction,
Naturwissenschaften
, p.(in press), (2014)
[20]. In some studies larger colonies produce more drones Factors determining the rearing and rejection of drones by the honeybee colony,
Anim. Behav.
, Volume 23, p.650-675, (1975)
[5] but see The effect of a plentiful supply of drone comb on colonies of honeybees,
J. Apic. Res.
, Volume 4, p.109-119, (1965)
[4]Protandrous reproduction in honey bees,
Envir. Entomol.
, Volume 10, p.359-361, (1981)
[6]. Number of drones produced in a colony can be determined genetically Observations sur la variabilité de la production du couvain de mâles dans les colonies d'abeilles (Apis mellifica L.),
Apidologie
, Volume 3, Number 4, p.291–307, (1972)
[21] and differs between subspecies Differential drone production by Africanized and European honey bee colonies,
Apidologie
, Volume 18, p.61-67, (1987)
[22]. Production of drones is smaller in presence of drone brood Factors determining the rearing and rejection of drones by the honeybee colony,
Anim. Behav.
, Volume 23, p.650-675, (1975)
[5] or adult drones Male reproductive parasitism: a factor in the africanization of European honey-bee populations.,
Science
, 1985 May 31, Volume 228, Issue 4703, p.1119-11121, (1985)
[23] but see Sperm utilization by honey bee queens: DNA fingerprinting analysis,
Applied Entomology and Zoology
, Volume 30, p.335-341, (1995)
[24].

There are suggestions that drones are costly to their colony Differences in drone and worker physiology in honeybees (Apis mellifera),
Apidologie
, Volume 36, Number 2, p.255-277, (2005)
[25], therefore, beekeepers often provide only worker foundation and remove combs with large number of drone cells. Colonies without drone combs produce more honey The effect of drone comb on a honey bee colony's production of honey,
Apidologie
, Volume 33, Number 1, p.75-86, (2002)
[26] but see Drone production in honey-bee colonies (Apis mellifera L.),
Nature
, Volume 199, p.789-790, (1963)
[16]The effect of a plentiful supply of drone comb on colonies of honeybees,
J. Apic. Res.
, Volume 4, p.109-119, (1965)
[4]Effects of drone comb on brood and honey production in honey bee colonies,
Ann. Entomol. Soc. Am.
, Volume 64, Number 4, p.954-956, (1971)
[27]Observations sur la variabilité de la production du couvain de mâles dans les colonies d'abeilles (Apis mellifica L.),
Apidologie
, Volume 3, Number 4, p.291–307, (1972)
[21]. It was not confirmed that colonies with smaller number of drone comb produce more workers Drone production in honey-bee colonies (Apis mellifera L.),
Nature
, Volume 199, p.789-790, (1963)
[16]The effect of a plentiful supply of drone comb on colonies of honeybees,
J. Apic. Res.
, Volume 4, p.109-119, (1965)
[4]Effects of drone comb on brood and honey production in honey bee colonies,
Ann. Entomol. Soc. Am.
, Volume 64, Number 4, p.954-956, (1971)
[27]. Although swarming colonies tend to rear more drones Protandrous reproduction in honey bees,
Envir. Entomol.
, Volume 10, p.359-361, (1981)
[6] the presence of drones does not affect tendency to swarm The effect of drone comb on a honey bee colony's production of honey,
Apidologie
, Volume 33, Number 1, p.75-86, (2002)
[26].

Queen prefers to lay eggs in worker cells than in drone cells The honeybee queen has the potential ability to regulate the primary sex ratio,
Appl Entomol Zool.
, Volume 31, p.247-254, (1996)
[28]. The proportion of eggs laid by the queen in drone cells was higher in spring than in summer The honeybee queen has the potential ability to regulate the primary sex ratio,
Appl Entomol Zool.
, Volume 31, p.247-254, (1996)
[28]. Queens who were forced to lay only drone eggs subsequently laid fewer drone eggs The honeybee queen influences the regulation of colony drone production,
Behavioral Ecology
, Volume 18, p.1092-1099, (2007)
[29]. This suggests that the queen can be involved in regulation of drone production. It was suggested that in colonies with older and unhealthy queens more drone cells are build Pheromones of social bees,
, London, p.218, (1987)
[30], however, this needs to be verified. Drone cell construction is not regulated by queen mandibular pheromones because removal of queens mandibular gland does not affect drone cell production The effect of mandibular gland extirpation on the longevity and attractiveness to workers of queen honey bees, Apis mellifera,
Annals of the Entomological Society of America
, Volume 57, p.73-74, (1964)
[31].

Theoretical models analysing possible proximate mechanisms of drone production control: Seasonal changes of drone numbers in a colony of the honeybee, Apis mellifera,
Ecological Modelling
, Volume 37, Number 3-4, p.155–166, (1987)
[32]Drone production in honeybee colonies: Controlled by a longlasting inhibitory pheromone from the drones?,
Journal of Theoretical Biology
, Volume 134, Number 3, p.309–318, (1988)
[33].
For review of drone production see Pheromones of social bees,
, London, p.218, (1987)
[30]Honeybee colony drone production and maintenance in accordance with environmental factors: an interplay of queen and worker decisions,
Insectes Sociaux
, Volume 57, Number 1, p.1-9, (2010)
[34].
Other references: The brood-rearing cycle of the honeybee,
Bull. U.S. Dep. Agric.
, Volume 1349, p.1-56, (1925)
[35]Seasonal brood rearing activity of the Cyprian honeybee,
J. Econ. Entomol.
, Volume 21, p.392-403, (1928)
[36]Untersuchungen über die Drohnenerzeugung im Bienenvolk,
Arch. Bienenkd.
, Volume 39, p.1–7, (1962)
[37]Drohnen im Natur-und Pflegevolk,
Allgemeine Deutsche Imkerzeitung
, Volume 39, p.100-105, (1969)
[38]The effect of induced changes in sexual asymmetry of honey bees (Apis mellifera) on swarming behaviour,
Biologia
, Volume 66, Number 3, p.535–542, (2011)
[39]Spatial distribution of honeybee drones Apis mellifera L. and certain factors affecting their production in the colony,
Department of Economic Entomology and Pesticides
, Cairo, Egypt, (2016)
[40].