Thursday, July 24, 2025

Burr Comb and Cross Comb in Langstroth Hives

Understanding Burr Comb and Cross Comb in Langstroth Hives

Burr Comb

Burr comb refers to small, irregular wax structures built by bees in unintended gaps within the hive, such as between frames, under the lid, or across the inner cover. These structures arise when spacing within the hive exceeds the ideal bee space, which is approximately 6 to 9 millimetres. In these situations, bees instinctively fill the space with wax or propolis (Bradbear, 2009; Seeley, 2010). Langstroth’s original design emphasized this precise spacing, noting that deviations would prompt bees to either block tight gaps with propolis or fill wide spaces with comb (Langstroth, 1853).

Cross Comb

Cross comb occurs when bees construct comb at angles that do not align with the frames. Instead of building straight within the plane of a frame, the comb stretches across adjacent frames. This results in disorganized construction that complicates inspections and often leads to the destruction of comb

during hive management (Winston, 1987).

Causes of Burr Comb and Cross Comb

Several factors contribute to the formation of burr and cross comb. First, improper bee space is a major culprit. If the space between hive components deviates from the 6–9 mm standard, bees will likely modify the space by constructing wax bridges or barriers. Misaligned or warped frames also create uneven gaps, encouraging the bees to build in unintended directions (Root, 1921).

Additionally, the absence of foundation or poorly attached foundation sheets can prompt bees to follow their instincts and create naturally oriented, and often irregular, combs (Hepburn & Radloff, 2011). Another significant cause is the orientation and leveling of the hive. Bees rely on gravity to construct vertical combs. If the hive is tilted, especially from side to side, the bees’ natural alignment is disrupted, leading to angled combs that result in cross combing (Seeley, 2010).

Furthermore, introducing swarms into boxes without drawn comb or foundation often results in haphazard comb building, particularly in deep boxes with no visual or structural guides (Crane, 1990).

Disadvantages of Burr Comb and Cross Comb

The disadvantages of burr and cross comb are numerous. They impede routine inspections by fusing frames together or obscuring frame edges, making it difficult to lift them without damaging bees or comb. This increases the risk of injuring the queen or destroying brood and honey stores. Additionally, broken burr comb filled with honey creates a sticky mess that attracts pests and disturbs the colony. Over time, irregular comb reduces usable hive space and complicates colony management (Delaplane, 2007).

Control and Prevention

To control and prevent burr and cross comb, several strategies can be implemented. Maintaining precise bee space is essential. This involves using quality equipment and regularly inspecting for warped or damaged frames. Providing bees with foundation sheets or comb starter strips guides proper comb construction, especially in new hives (Hepburn & Radloff, 2011).

Ensuring that hives are level, particularly from side to side, supports natural vertical comb alignment. Frequent early inspections during comb-building phases allow beekeepers to intervene before cross comb becomes extensive. In addition, using the correct number of frames to avoid excessive gaps in supers can prevent bees from constructing comb in unintended spaces.

Corrective actions include trimming irregular comb early and realigning frames. If done promptly, bees often adapt and reconstruct comb in the desired direction. Consistent management and attention to structural detail ensure that Langstroth hives function as intended, reducing disruptions and supporting colony health.

Raising Queens from Swarm, Supersedure, and Emergency Cells

In managed beekeeping, the type of queen cell used to raise a new queen can significantly influence the overall success and productivity of the resulting colony. Beekeepers commonly rely on swarm, supersedure, or emergency queen cells when rearing replacement queens or expanding colonies. However, these cell types vary in terms of biological origin, the circumstances in which they are produced, and the quality of queens they generate. Drawing from contemporary scientific literature, this article explores these differences in depth and outlines when and why each queen cell type might be preferred.

Queen Cell Types and Their Biological Origins

Swarm cells are produced by colonies in preparation for swarming—a natural method of colony reproduction. These elongated, peanut-shaped cells are typically located along the edges of brood comb and are created under conditions of resource abundance and strong colony vigor. The queen that lays the eggs destined to become swarm queens is often in good reproductive condition, and the resulting larvae are well-fed by nurse bees throughout their development (Simone-Finstrom et al., 2016). Because they are raised under planned and favorable circumstances, swarm cells tend to yield robust queens.

Supersedure cells, in contrast, are constructed when the colony determines that the reigning queen is no longer performing optimally—often due to declining pheromone output or reduced egg-laying capacity. These cells are usually found in the central part of the comb. Unlike swarm situations, the colony remains queenright during supersedure events. The objective here is not reproduction but internal queen replacement to sustain colony performance. While the environmental conditions are often stable, the urgency for replacement is typically lower than in emergency scenarios (Winston, 1987).

Emergency cells arise when a colony loses its queen abruptly—either through beekeeper error, predation, or disease. In such cases, the colony responds by identifying young worker larvae, less than three days old, and rearing them into queens by feeding them royal jelly and enclosing them in specially extended vertical cells. This rapid response mechanism ensures the survival of the colony, but it imposes developmental constraints on the replacement queen due to time limitations and suboptimal nutritional conditions (Tarpy et al., 2000).

Scientific Insights into Queen Quality

Research indicates that queens reared under swarm conditions consistently outperform those produced under supersedure or emergency conditions. Swarm queens tend to be larger in size, exhibit more developed reproductive organs, and have higher mating success rates. This superiority is attributed to the favorable colony status and abundant resources available during their rearing period (Gilley, 2001).

Supersedure queens are generally acceptable, though studies show a degree of variability in their reproductive fitness depending on the age and condition of the queen they replace and the overall health of the colony (Hatch et al., 1999). Emergency queens are more likely to be of lower quality. Since they are derived from worker-destined larvae and raised hastily, they may be smaller, less fertile, and more susceptible to mating failures or early supersedure (Tarpy et al., 2000).

Guidance for Beekeepers: Choosing the Right Cell Type

Swarm cells are best suited for planned colony divisions, such as raising nucleus colonies (nucs) or executing controlled expansions. These cells tend to yield queens of excellent quality, but they also require close monitoring to avoid uncontrolled swarming.

Supersedure cells are most appropriate for replacing a failing queen within an otherwise stable colony. While these queens are not as reliably robust as swarm queens, they are often sufficient for colony maintenance, especially if the workers make the replacement decision gradually and under stable environmental conditions.

Emergency queen cells should be reserved for true crises—situations in which the colony is suddenly queenless. While they serve a critical survival function, the resulting queens are often inferior in terms of reproductive performance and longevity. In such cases, beekeepers may consider introducing a purchased or grafted queen instead, to avoid compounding colony stress with a suboptimal queen.

Considerations Affecting Queen Success

Several additional factors influence the success of queens reared from any of these cell types. First, timing is crucial. Queens raised from swarm and supersedure cells develop over the standard 16-day period from egg to emergence. In contrast, emergency queens often develop from slightly older larvae, compressing their developmental window and potentially compromising physical development (Winston, 1987).

Second, nutritional conditions play a fundamental role in queen quality. Regardless of the cell type, inadequate feeding of larvae during early development will result in smaller, less capable queens. Even well-situated swarm cells can produce poor queens if the colony is experiencing nutritional stress.

Finally, successful mating is essential for a viable queen. Factors such as weather, availability of drones, and mating flight success all influence whether a seemingly well-developed queen will establish a strong laying pattern. A queen’s quality is not solely defined by her physical characteristics but also by her ability to mate effectively and begin laying fertilized eggs promptly (Delaney et al., 2009).

References

Delaney, D. A., Keller, J. J., Caren, J. R., & Tarpy, D. R. (2009). The physical, insemination, and reproductive quality of honey bee queens (Apis mellifera). Apidologie, 40(5), 563–572. https://doi.org/10.1051/apido/2009049

Gilley, D. C. (2001). The behavior of honey bees (Apis mellifera ligustica) during swarm cell construction. Animal Behaviour, 61(1), 1–9. https://doi.org/10.1006/anbe.2000.1552

Hatch, S., Tarpy, D. R., & Fletcher, D. J. C. (1999). Worker regulation of emergency queen rearing in honey bee colonies and the resultant variation in queen quality. Insectes Sociaux, 46, 372–377. https://doi.org/10.1007/s000400050158

Simone-Finstrom, M., Li-Byarlay, H., Huang, M. H., Strand, M. K., Rueppell, O., & Tarpy, D. R. (2016). Migratory management and environmental conditions affect lifespan and oxidative stress in honey bees. Scientific Reports, 6, 32023. https://doi.org/10.1038/srep32023

Tarpy, D. R., Hatch, S., & Fletcher, D. J. C. (2000). The influence of queen age and quality during queen replacement in honeybee colonies. Animal Behaviour, 59(1), 97–101. https://doi.org/10.1006/anbe.1999.1278

Winston, M. L. (1987). The Biology of the Honey Bee. Cambridge, MA: Harvard University Press.

Tuesday, July 22, 2025

Historical Origin of the Langstroth Hive

Beekeeping Before Langstroth

Before the 1850s, beekeepers commonly used fixed comb hives such as straw skeps or hollow log gums. In these traditional hives, bees attached their comb directly to the internal surfaces, making it impossible to harvest honey or inspect brood without tearing apart the comb (Crane, 1999). Honey harvesting typically resulted in the destruction of the colony, often by suffocating the bees using smoke or sulfur (Bradbear, 2009). These limitations made traditional beekeeping unsustainable and inefficient.

Discovery of the Bee Space (1851)

In 1851, Reverend Lorenzo Lorraine Langstroth, an American clergyman and beekeeper, observed that when a space of approximately 6 to 9 millimeters was maintained between combs or hive parts, bees neither built comb in the gap nor sealed it with propolis. He termed this discovery the “bee space” (Langstroth, 1853). Langstroth then designed a rectangular hive using removable wooden frames suspended inside a box, with all gaps spaced precisely to preserve the bee space. This innovation allowed frames to be removed for inspection or honey harvesting without destroying combs or aggravating the bees (Root, 1918).

Patent of the Movable Frame Hive (1852)

Langstroth patented his movable frame beehive on October 5, 1852, under U.S. Patent No. 9300. The design incorporated internal spacing that respected the bee space, allowing beekeepers to manage colonies with unprecedented ease and efficiency (Graham, 1992). By the end of that year, Langstroth had operationalized over 100 of these hives in his apiary. The design made it possible to harvest honey without killing bees or destroying wax combs, representing a major advancement in the field (Crane, 1999).

Publication and Impact (1853)

In 1853, Langstroth published A Practical Treatise on the Hive and Honey-Bee, a comprehensive manual explaining his beekeeping method and hive design (Langstroth, 1853). The book quickly became a foundational text for American beekeepers, offering detailed guidance on colony management, seasonal care, and honey extraction. The principles of movable frames, modular hive construction, and bee space became standard features in modern hive design.

Global Influence and Legacy

Although European beekeepers like Jan Dzierżon and August von Berlepsch had explored similar concepts earlier, Langstroth was the first to fully implement the bee space principle in a practical and replicable hive system (Crane, 1999). His invention transformed beekeeping from a destructive to a sustainable practice and earned him recognition as the father of American beekeeping (Root, 1918). The Langstroth hive quickly spread internationally and remains the most widely used hive today, forming the basis for approximately 75 percent of the world’s managed hives (Adjare, 1990; Bradbear, 2009).

References

Adjare, S. O. (1990). Beekeeping in Africa. Food and Agriculture Organization of the United Nations.

Bradbear, N. (2009). Bees and their role in forest livelihoods: A guide to the services provided by bees and the sustainable harvesting, processing and marketing of their products. Food and Agriculture Organization of the United Nations.

Crane, E. (1999). The world history of beekeeping and honey hunting. Routledge.

Graham, J. M. (Ed.). (1992). The hive and the honey bee. Dadant & Sons.

Langstroth, L. L. (1853). A practical treatise on the hive and honey-bee. Hopkins, Bridgman & Company. https://www.gutenberg.org/ebooks/24583

Root, A. I. (1918). The ABC and XYZ of bee culture. A. I. Root Company.

U.S. Patent and Trademark Office. (1852). Movable-frame beehive: Patent No. 9300. Filed October 5, 1852.

📚 View Full References

Adjare, S. O. (1990). Beekeeping in Africa. FAO.
Bradbear, N. (2009). Bees and their role in forest livelihoods. FAO.
Crane, E. (1999). The world history of beekeeping and honey hunting. Routledge.
Graham, J. M. (Ed.). (1992). The hive and the honey bee. Dadant & Sons.
Langstroth, L. L. (1853). A practical treatise on the hive and honey-bee. Gutenberg. [Link]
Root, A. I. (1918). The ABC and XYZ of bee culture. A. I. Root Co.
U.S. Patent and Trademark Office (1852). Patent No. 9300.

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Monday, July 21, 2025

Why Do Drone Bees Die After Mating?

Drone honey bees (Apis mellifera) are biologically specialized for one function: mating with a queen. Unlike worker bees, drones do not forage, care for the brood, or build comb. They exist solely to pass on genetic material. However, this act comes at a fatal cost. A drone that successfully mates dies immediately afterward. This article explains the biological mechanism and evolutionary reasoning behind drone mortality after mating.

Drone Congregation Areas: Where Mating Begins

Honey bee mating does not occur randomly. Instead, it takes place at designated locations called Drone Congregation Areas (DCAs). These are stable open air zones typically found 10 to 40 meters above ground level. They often form in open spaces near natural features such as forest edges, tree gaps, or clearings (Koeniger, Koeniger, & Tingek, 2005).

Thousands of drones from different colonies gather daily in DCAs during mating season. Virgin queens enter these areas during their nuptial flights and are pursued midair by drones. Mating occurs in flight, and a queen may visit several DCAs, typically mating with 12 to 20 drones over one or more days (Baudry et al., 1998). This system encourages genetic mixing and reduces the risk of inbreeding.

The Mating Process and Drone Physiology

When a drone succeeds in mating, he uses an internal structure called the endophallus to inseminate the queen. This organ is everted, or turned inside out, under intense pressure during ejaculation. A single ejaculation can transfer up to 90 million sperm cells (Koeniger et al., 2014).

The force of this transfer is so extreme that the endophallus ruptures and becomes detached inside the queen’s reproductive tract, forming a temporary mating plug. As a result of this rupture, the drone experiences fatal abdominal damage, including tearing of tissues and hemolymph loss. Death follows within seconds (Woyke, 1958).

Why Drones Die After Mating

Drone mortality is not accidental. It is a direct result of their anatomical specialization. Unlike males in many insect species who survive and can mate multiple times, honey bee drones are structured for single use reproduction. The mating event causes irreversible physical damage, making post mating survival biologically impossible.

Evolutionary Significance

From an evolutionary standpoint, this reproductive strategy represents a terminal investment. The drone maximizes his reproductive success through one complete mating. The detachment of the endophallus ensures full sperm transfer and may briefly deter further mating attempts by other drones (Baer, 2005).

This seemingly costly strategy benefits the colony. Queens store the sperm of all mates in a structure called the spermatheca, using it to fertilize eggs throughout their lives. By mating with multiple drones, the queen ensures genetic diversity and resilience within the colony.

References

Baer, B. (2005). Sexual selection in Apis bees. Apidologie, 36(2), 187–200. https://doi.org/10.1051/apido:2005012

Baudry, E., Solignac, M., Garnery, L., Gries, M., Cornuet, J. M., & Koeniger, N. (1998). Relatedness among drone congregations of the honeybee (Apis mellifera L.). Proceedings of the Royal Society B: Biological Sciences, 265(1392), 2009–2014. https://doi.org/10.1098/rspb.1998.0533

Koeniger, G., Koeniger, N., & Tingek, S. (2005). Mating Biology of Honey Bees (Apis). International Bee Research Association.

Koeniger, G., Koeniger, N., Ellis, J., & Connor, L. J. (2014). Queen Mating and Reproduction in Honey Bee Colonies. Wicwas Press.

Woyke, J. (1958). Natural and artificial insemination of queen honeybees. Bee World, 39(3), 57–65. https://doi.org/10.1080/0005772X.1958.11095037

Sources and Further Reading

You can view or access the sources cited in this article below:

Title	Authors / Source	Link
Sexual selection in Apis bees	Baer (2005) – Apidologie	View Article
Relatedness among drone congregations of the honeybee	Baudry et al. (1998) – Proceedings of the Royal Society B	View Article
Mating Biology of Honey Bees (Apis)	Koeniger, Koeniger, & Tingek (2005) – IBRA	Publisher Page
Queen Mating and Reproduction in Honey Bee Colonies	Koeniger et al. (2014) – Wicwas Press	View Book
Natural and artificial insemination of queen honeybees	Woyke (1958) – Bee World	View Article

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Sunday, July 20, 2025

Do Bees Eat Honey?

Do Bees Eat Honey?

Yes, honey bees (Apis mellifera) eat the honey they produce. Honey is their primary source of carbohydrates and is vital not only for daily energy but also for other metabolic functions such as thermoregulation, brood care, and wax secretion (Quinlan et al., 2023; Huang, 2018).

Nutritional Function of Honey

Honey is composed primarily of simple sugars—fructose and glucose—making up about 80–85% of its content, with water accounting for 15–17% (Martinotti & Ranzato, 2023). It also contains trace amounts of enzymes, amino acids, vitamins, and minerals (Rao et al., 2016). This sugar-rich composition makes honey an efficient energy source. Worker bees consume honey to fuel flight, fanning, foraging, and maintaining hive temperature. During cold periods or nectar scarcity, bees rely entirely on their honey reserves for survival (Mississippi State University Extension, 2019).

Honey and Wax Production

Beyond energy, honey is also essential in wax production. Worker bees consume large amounts of honey to generate wax scales from abdominal glands. According to Hepburn (1986), it takes approximately 6.6 to 8.4 kg of honey to produce 1 kg of beeswax, highlighting the metabolic cost of comb building. The wax is critical for constructing brood cells and storage areas for pollen and honey.

Honey Consumption in Tropical Climates

In tropical regions such as sub-Saharan Africa where winter is absent, honey bees do not enter dormancy but still rely on stored honey during seasonal nectar dearths. For example, during long dry spells or periods of intense heat when flowering plants are sparse, bees reduce foraging and consume stored honey to maintain basic colony functions (Adjare, 1990; Nuru et al., 2015). These seasonal shortages mimic winter scarcity in temperate zones in terms of resource stress.

Do Bees Eat Honey Before Rain or If Harvest Is Delayed?

Yes. Bees can detect atmospheric changes, including humidity and pressure drops that signal impending rain. In anticipation, they increase feeding on stored honey to prepare for confinement during rainy days (Crane, 1990). Similarly, if honey remains unharvested and environmental conditions change—such as onset of a rainy season—bees may consume surplus honey, especially if foraging is curtailed. In such cases, beekeepers may find reduced yields if harvest is delayed.

Furthermore, the timing of harvesting is crucial. Delayed harvesting during or after peak nectar flow can coincide with the colony shifting to consumption rather than storage—especially if pollen becomes limited or nectar stops flowing. A study by Fichtl & Admasu (1994) in Ethiopia noted that local beekeepers often lose part of the honey yield if not collected early, as bees consume the surplus during late dry seasons.

How Much Honey Do Bees Consume?

An adult worker bee consumes approximately 11 mg of honey daily for metabolic needs (Huang, 2018). A standard colony of 50,000 bees may consume up to 1 kg of honey per day under active foraging or brood-rearing conditions. For long-term survival during scarcity periods, colonies may require 30–45 kg of honey reserves, depending on region and hive size (Mississippi State University Extension, 2019).

Sources and Downloads

You can view or download the academic and institutional sources cited in this article below:

Title	Authors / Source	Download / View
Carbohydrate Nutrition & Overwintering Bees	Quinlan et al. (2023) – Journal of Insect Science	View Article
Feeding Honey Bees	Huang (2018) – MSU Extension	Download PDF
Colony Growth & Seasonal Management	Mississippi State University Extension (2019)	Download PDF
Beehive Products and Healing	Martinotti & Ranzato (2023) – Cosmetics	View Article
Therapeutic Effects of Honey	Rao et al. (2016) – Brazilian Journal of Pharmacognosy	View Article
Honeybee Wax Biology	Hepburn (1986) – Bee World	View Article
Beekeeping in Africa	Adjare (1990) – FAO	View Full Book
Honeybee Flora of Ethiopia	Fichtl & Admasu (1994) – GTZ	Download PDF
Bees and Beekeeping: World Resources	Crane (1990) – Book Excerpt	View Book Online

References

Adjare, S. O. (1990). Beekeeping in Africa. FAO Agricultural Services Bulletin 68/6. Food and Agriculture Organization of the United Nations.

Crane, E. (1990). Bees and Beekeeping: Science, Practice and World Resources. Heinemann Newnes.

Fichtl, R., & Admasu, A. (1994). Honeybee flora of Ethiopia. Deutsche Gesellschaft für Technische Zusammenarbeit (GTZ).

Hepburn, H. R. (1986). Honeybee wax: An overview of its biology, collection, properties and functions. Bee World, 67(3), 119–132. https://doi.org/10.1080/0005772X.1986.11098954

Huang, Z. (2018). Feeding honey bees (Bulletin E-3369). Michigan State University Extension.

Martinotti, S., & Ranzato, E. (2023). Applications of beehive products for wound repair and skin care. Cosmetics, 10(5), 127. https://doi.org/10.3390/cosmetics10050127

Mississippi State University Extension. (2019). Colony growth and seasonal management of honey bees. https://extension.msstate.edu/sites/default/files/publications/publications/P3052_web.pdf

Quinlan, G., Winge, P., Medici, S., & Hood, W. M. (2023). Carbohydrate nutrition associated with health of overwintering honey bees. Journal of Insect Science, 23(6), 16. https://doi.org/10.1093/jisesa/iead079

Rao, P. V., Krishnan, K. T., Salleh, N., & Gan, S. H. (2016). Biological and therapeutic effects of honey produced by honey bees and stingless bees: A comparative review. Brazilian Journal of Pharmacognosy, 26(5), 657–664. https://doi.org/10.1016/j.bjp.2016.03.011

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Thursday, July 17, 2025

Better Queens

Successful Queen Rearing

Queen rearing is a vital technique in apiculture that enables beekeepers to propagate desirable traits, maintain colony health, and increase productivity. To achieve consistent success, queen rearing must be based on a strong understanding of honey bee biology and managed under optimal environmental and technical conditions. This article outlines the essential prerequisites and considerations for effective queen rearing, backed by peer-reviewed scientific sources.

Use of Strong and Disease-Free Colonies

A strong cell builder colony is the backbone of any queen rearing program. It must be populous, rich in nurse bees, and free of diseases like American foulbrood (Paenibacillus larvae) and Varroa destructor infestations. Healthy nurse bees are critical for producing copious amounts of royal jelly required for developing quality queens (Delaplane et al., 2013).

Selection of High-Quality Breeder Stock

The genetic foundation of any queen starts with the selection of superior breeder queens and drones. Traits such as high honey production, hygienic behavior, gentleness, and disease resistance should guide selection. As highlighted by Tarpy et al. (2000), queen mating success and colony genetic diversity significantly influence colony performance and resilience.

Abundant and Mature Drone Populations

Mating success depends heavily on the availability of mature drones. Honey bee queens typically mate with an average of 14 drones, with ranges from 10 to over 20, during several mating flights within 5–10 days post-emergence (Delaney et al., 2011; Koeniger et al., 2005). Drone congregation areas (DCAs) often include thousands of drones from numerous colonies, promoting genetic diversity (Tarpy & Page, 2000).

Favorable Environmental Conditions

Successful queen rearing is seasonal and climate-dependent. Ideal periods are during spring and early summer, when colonies are naturally expanding and floral resources are abundant. Queens require clear, warm days (typically above 21°C) for successful mating flights (Koeniger et al., 2005). Poor weather can delay mating or reduce mating quality.

Accurate Grafting and Larval Selection

Larvae should be grafted between 12 and 24 hours old, before worker fate is irreversible. Laidlaw and Page (1997) demonstrated that older larvae result in queens of lower quality, with reduced pheromone output and reproductive capacity. Tools such as grafting pens or Jenter systems aid in transferring larvae safely.

Proper Feeding and Nutrition

Feeding colonies with sugar syrup and protein supplements enhances royal jelly production and supports larval health. Nutritional stress reduces the quality of the queen. Brodschneider and Crailsheim (2010) emphasized the role of balanced nutrition in colony development and immune strength.

Well-Managed Mating Nuclei

Mating nuclei must be adequately stocked with young bees, food reserves, and must remain queenless before cell introduction. Placement near DCAs and adequate spacing of nuclei prevent confusion during queen return flights and maximize mating success (Koeniger et al., 2005).

Monitoring and Record Keeping

Accurate records on grafting success, queen emergence, and mating results are crucial. Tracking lineage, colony behavior, and productivity over time allows for continual improvement in queen quality (Delaney et al., 2009).

References

Brodschneider, R. and Crailsheim, K. (2010) 'Nutrition and health in honey bees', Apidologie, 41(3), pp. 278–294. https://doi.org/10.1051/apido/2010012

Delaney, D.A., Keller, J.J., Caren, J.R. and Tarpy, D.R. (2009) 'The physical, insemination, and reproductive quality of honey bee queens (Apis mellifera)', Apidologie, 42(1), pp. 1–13. https://doi.org/10.1051/apido/2009049

Delaplane, K.S., van der Steen, J. and Guzman-Novoa, E. (2013) 'Standard methods for estimating strength parameters of Apis mellifera colonies', Journal of Apicultural Research, 52(1), pp. 1–12. https://doi.org/10.3896/IBRA.1.52.1.03

Koeniger, N., Koeniger, G. and Pechhacker, H. (2005) 'The nearer the better? Drones (Apis mellifera) prefer nearer drone congregation areas', Apidologie, 36(4), pp. 413–420.

Laidlaw, H.H. and Page, R.E. (1997) Queen Rearing and Bee Breeding. Cheshire, CT: Wicwas Press.

Tarpy, D.R., Nielsen, R. and Nielsen, D.I. (2000) 'A scientific note on the number of drone matings in natural honey bee populations', Apidologie, 31(3), pp. 343–345. https://doi.org/10.1051/apido:2000126

Tarpy, D.R. and Page, R.E. (2000) 'No behavioral control over mating frequency in queen honey bees (Apis mellifera L.)', Animal Behaviour, 59(4), pp. 875–882. https://doi.org/10.1006/anbe.1999.1386

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