Bee Breeding & Selection: Choosing for Traits That Matter

Every colony in your apiary carries a genetic blueprint that determines how it behaves, how much honey it produces, how well it survives winter, and how effectively it resists disease. Most beekeepers treat genetics as something that arrives with a package or a queen in the mail — an input they have little control over. But honey bee breeding gives you the ability to shape the future of your bees, one generation at a time.

This guide covers the science and practice of honey bee breeding and selection. It is written for beekeepers who have at least a few seasons of experience, who understand basic colony management, and who are ready to think deliberately about the genetic quality of their stock. Whether you want gentler bees, better mite resistance, or colonies that thrive in your specific climate, the principles here will give you a framework for making real progress.

Why Breed Your Own Bees

Genetic Improvement Over Time

Package bees and commercially available queens come from operations that breed for broad market appeal — shipping hardiness, early buildup, and general reliability. These are worthwhile traits, but they are not necessarily the traits that matter most in your apiary, in your climate, with your management style.

When you select your own breeder queens from colonies that perform well under your specific conditions, you concentrate favorable alleles with every generation. Over three to five years of deliberate selection, you can produce stock that meaningfully outperforms the generic alternatives in your operation.

Local Adaptation

Honey bees in subtropical South Florida face pressure from Small Hive Beetles, high humidity, and year-round brood cycles. Bees in northern Minnesota contend with five months of confinement, dramatic temperature swings, and short foraging seasons. No single strain excels in both environments.

Breeding from your own survivor stock — colonies that have already proven they can handle your local challenges — produces bees that are progressively better adapted to your region. This is not a controversial idea; it is the foundation of livestock improvement in every agricultural species. Bees are no different.

Reducing Dependency

Commercial queen producers face their own challenges: shipping losses, disease pressure in large-scale operations, and the logistical difficulty of producing queens on demand across a wide geographic range. By developing your own queen-rearing and breeding skills, you reduce your dependence on external suppliers and gain flexibility in how you manage your apiary.

🧬 Key Insight: Breeding is not about creating the "perfect bee." It is about incrementally shifting the average performance of your apiary in the direction you choose. Small, consistent gains compound over time.

Honey Bee Genetics Basics

Understanding a few core principles of honey bee genetics is essential before making breeding decisions. Honey bees have a reproductive system that differs fundamentally from mammals and most other livestock.

Haplodiploidy

Honey bees are haplodiploid. Females (workers and queens) develop from fertilized eggs and are diploid — they carry two sets of chromosomes, one from each parent. Males (drones) develop from unfertilized eggs and are haploid — they carry only one set of chromosomes, inherited entirely from their mother.

This has profound implications for breeding:

1. Drones express all of their mother's genes. A drone has no father. Every allele in his genome came directly from the queen that produced him. This means a drone fully expresses his mother's genetic potential — there are no hidden recessive traits.
2. Workers share more genes with each other than with their own offspring. Full-sister workers share, on average, 75% of their genes (because they share 100% of the paternal contribution from a single drone father and, on average, 50% of the maternal contribution). This relatedness is the genetic foundation for eusocial behavior.
3. The queen's mating partners matter enormously. Because a queen mates with multiple drones, the workers in a single colony belong to multiple patrilines — subfamilies that share the same mother but different fathers. This genetic diversity within the colony affects everything from disease resistance to foraging efficiency.

Polyandry

A queen typically mates with 12 to 20 drones during her nuptial flights. This extreme polyandry is not accidental — it is an evolutionary strategy that increases genetic diversity within the colony. A colony with high patriline diversity is more resistant to disease, shows more consistent foraging behavior across changing conditions, and maintains more stable thermoregulation.

From a breeding perspective, polyandry means that even if you have an outstanding breeder queen, her daughters will not be uniform. They will carry genes from multiple drone fathers, some desirable and some less so. This is why repeated selection across generations is necessary — you cannot fix traits in a single generation.

Subspecies and Geographic Races

Apis mellifera is native to Europe, Africa, and western Asia. Over millennia, geographic isolation produced distinct subspecies (often called "races") adapted to different climates and floral patterns. The major subspecies relevant to North American beekeeping include:

Most commercial queens are not pure subspecies but admixed populations with ancestry from multiple races. This is not inherently bad — hybrid vigor is real — but it does mean you cannot reliably predict colony traits from subspecies labels alone. Performance evaluation of individual colonies matters more than the name on the queen package.

Traits Worth Selecting For

Not all traits are equally important, and some are easier to measure than others. The table below summarizes the major traits that beekeepers select for, along with practical considerations for each.

Trait Prioritization

You cannot optimize for every trait simultaneously. Trade-offs exist:

• High honey production sometimes correlates with higher swarm tendency.
• Strong Varroa resistance (VSH lines) may produce slightly less honey than non-resistant stock in the absence of mite pressure.
• Extreme gentleness can sometimes come at the cost of reduced defensiveness against robbing and predators.

The most successful breeding programs focus on two or three primary traits and accept good-enough performance in the others. For most beekeepers in 2026, the recommended priority order is:

1. Varroa resistance — Without this, nothing else matters. Mites are the number one cause of colony loss.
2. Winter survival — Dead colonies produce no honey.
3. Gentleness — Bees you are afraid to inspect get neglected.
4. Honey production — Important, but secondary to survival.

⚠️ Reality Check: Heritability estimates for honey bee traits are approximate and vary by population. Do not expect rapid, dramatic change. Meaningful genetic progress in honey bees typically takes 5–10 years of consistent selection. Patience is not optional — it is the process.

Instrumental Insemination

What It Is

Instrumental insemination (II) is a technique in which a queen is artificially inseminated with collected drone semen using specialized equipment — a syringe, a hook, and a stereomicroscope. Developed in the 1920s and refined over subsequent decades, II gives the beekeeper complete control over the genetic contribution of both parents.

When to Use It

II is appropriate in several specific situations:

1. Controlled breeding programs where you need to know the exact sire and dam of each queen — for example, crossing a VSH queen with drones from a high-production line to evaluate hybrid performance.
2. Maintaining pure lines of selected stock without contamination from uncontrolled drone sources.
3. Research settings where experimental rigor requires known parentage.

Limitations

II is not a practical tool for most beekeepers:

• Equipment cost — A quality II setup costs $2,000–$4,000 (microscope, syringe, needles, CO2 delivery system).
• Skill requirement — Achieving consistent, high-quality inseminations requires extensive practice. Expect to practice on 100+ queens before achieving reliable results.
• Reduced genetic diversity — II queens typically receive semen from a small number of drones (often 8–12), which is far fewer than natural mating (12–20+). This limits the patriline diversity within the resulting colony.
• Semen collection logistics — Drone semen must be collected fresh and used promptly. Long-term storage techniques (cryopreservation) exist but are not yet practical for routine use.
• No guarantee of superiority — II controls parentage, not outcome. A poorly chosen cross can produce mediocre stock just as easily as a well-chosen one.

A Balanced View

For most small-to-mid-scale beekeepers, the time and money invested in mastering II would produce better returns if spent on improving evaluation skills, increasing colony numbers (and thus the pool from which to select), or implementing a rigorous open-mating breeding program. II is a powerful tool for specialists and researchers, but it is not a prerequisite for meaningful genetic improvement.

Open Mating & Drone Saturation

How Open Mating Works

In open mating, virgin queens fly to drone congregation areas (DCAs) — traditional flyway sites where hundreds or thousands of drones from surrounding colonies gather. A queen typically makes two or three mating flights over several days, mating with multiple drones at the DCA each time.

The beekeeper controls the maternal genetics (by choosing which queens to rear virgins from) and can partially influence the paternal genetics (by flooding the area with drones from selected colonies), but cannot guarantee the exact drone composition of the mating.

Drone Saturation: The Practical Approach

Drone saturation is the strategy of producing large numbers of drones from your best colonies in the area where your virgins will mate. The goal is to tilt the odds: even though you cannot prevent uncontrolled drones from mating with your queens, you can ensure that a significant proportion of the sperm stored by each queen comes from your selected lines.

Steps for an effective drone saturation program:

1. Identify your top 3–5 colonies based on your trait evaluation scores.
2. Insert drone foundation into these colonies 40–50 days before you plan to have virgin queens ready to mate. Drones take 24 days to develop and need another 10–14 days to reach sexual maturity.
3. Limit drone production in average and below-average colonies by using worker-sized foundation and cutting out any drone comb they build.
4. Time your queen rearing so that virgin queens emerge when your selected drones are flying in peak numbers.
5. Locate mating nucs within half a mile of your drone-producing colonies, if possible. Queens typically mate within a mile of their hive, and drones typically fly to DCAs within 2 miles of their home colony.

Realistic Expectations

Even with aggressive drone saturation, you should expect that 30–50% of the drones at any DCA will come from colonies outside your control (unless your apiary is extremely isolated). This means open-mated queens will always carry some unselected genetics.

This is not a reason to avoid open mating. Over multiple generations, consistent drone saturation and careful selection of breeder queens will shift the genetic composition of your local drone population. Many of the most successful breeding programs in the world — including programs that have produced genuinely mite-resistant stock — rely primarily on open mating.

🐝 Practical Tip: If you have a neighbor within 2 miles who keeps bees, talk to them about your breeding goals. Collaborative drone saturation — where multiple beekeepers in an area agree to produce drones from their best stock — dramatically improves the effectiveness of open-mating programs. Cooperative breeding is one of the most underutilized strategies in beekeeping.

VSH (Varroa Sensitive Hygiene) Breeding

Understanding VSH

Varroa Sensitive Hygiene (VSH) is a specific form of hygienic behavior in which worker bees detect and remove brood cells that are infested with reproducing Varroa mites. It is distinct from general hygienic behavior (removing dead or diseased brood) in its specificity: VSH bees target cells where mites are actively reproducing, interrupting the mite reproductive cycle.

The VSH trait was first identified by researchers at the USDA Honey Bee Breeding, Genetics, and Physiology Laboratory in Baton Rouge, Louisiana, in the 1990s. It has since been confirmed as one of the most effective natural mechanisms for Varroa population control.

How VSH Works Mechanically

1. Detection — VSH workers detect the chemical signals (likely changes in cuticular hydrocarbons or volatile compounds) produced by brood cells containing reproducing mites.
2. Uncapping — The workers uncap the infested cell.
3. Removal — The workers remove the pupa along with the foundress mite and any daughter mites she has produced.
4. Interruption — By removing the pupa before daughter mites have matured and mated, the workers prevent those mites from infesting additional cells.

A colony with strong VSH expression can reduce Varroa reproduction by 50–80%, dramatically slowing the growth of the mite population and reducing or eliminating the need for chemical treatments.

Selecting for VSH

There are two primary approaches to selecting for VSH:

1. Field Selection Based on Mite Counts

This is the most practical approach for most beekeepers:

• Perform monthly alcohol wash mite counts on all colonies.
• Identify colonies that consistently show mite counts below the treatment threshold (typically 2–3 mites per 300 bees) even without treatment.
• Use these colonies as your breeder queens.
• Over time, the frequency of VSH genes in your population will increase.

The limitation of this approach is that low mite counts can result from factors other than VSH — for example, grooming behavior, short brood cycles, or simply a recent treatment. It is a proxy, not a direct measurement.

2. Direct VSH Assay

For more precise selection:

1. Prepare a frame of brood with known Varroa infestation (either by introducing mites into cells or by using naturally infested brood).
2. Place the frame in the test colony.
3. After 24–48 hours, examine the frame and record the percentage of infested cells that have been uncapped and the brood removed.
4. Colonies that remove more than 80% of infested cells within 48 hours show strong VSH expression.

This method is more labor-intensive but provides direct evidence of the VSH trait.

VSH Stock Sources

Several queen producers in the United States specialize in VSH stock or stock with significant VSH expression:

• USDA Baton Rouge VSH lines — The original source material; available through authorized distributors.
• Minnesota Hygienic — Developed at the University of Minnesota; strong general hygiene with some VSH expression.
• Puget Sound Bees — VSH-selected stock adapted to the Pacific Northwest.
• Various regional breeders — Many local queen producers now incorporate VSH genetics into their lines.

Introducing VSH queens into your apiary and then selecting daughter queens from those that perform best in your conditions is a practical way to build Varroa resistance into your stock.

Russian & Other Resistant Lines

Russian Honey Bees

Russian honey bees (A. m. caucasia region origin, not literally from Russia today) were imported to the United States through a USDA program in the late 1990s and early 2000s. They originated from the Primorsky Krai region of far eastern Russia, where Varroa destructor has been present for over a century — meaning these bees have been under natural selection for mite resistance for far longer than any Western stock.

Characteristics of Russian bees:

• Strong Varroa resistance — They express VSH behavior, grooming behavior, and other mite-fighting traits at high frequency.
• Conditional brood production — They ramp up brood rearing quickly when forage is available and slow down during dearths. This is energy-efficient but can catch beekeepers off guard if they expect the steady brood production typical of Italian stock.
• Moderate honey production — Generally comparable to Italian stock, sometimes higher in areas with strong late-season flows.
• Slightly more defensive — Not hot by any means, but less placid than typical Italian bees. Most beekeepers find them perfectly manageable.
• Strong swarming tendency — This is a genuine management challenge. Russian colonies require attentive swarm prevention.

Other Notable Resistant Lines

A Critical Point on Stock Selection

No stock is a silver bullet. Russian bees can still suffer mite damage if the mite load overwhelms their natural defenses. Saskatraz bees can still die in a harsh winter if they are not managed properly. Stock selection is one component of a comprehensive management strategy — it does not replace monitoring, treatment when necessary, and good husbandry.

The strongest long-term strategy is to introduce genetics from multiple resistant sources into your apiary and then select from the resulting population for the individuals that perform best under your specific conditions. This combines the benefits of existing breeding programs with the power of local adaptation.

Local Adaptation & Survivor Stock

The Case for Survivor Stock

The most compelling data in honey bee breeding comes not from controlled studies but from a simple observation: in any apiary that has been established for several years, some colonies die and some survive, even when managed identically. The survivors carry genes that helped them handle whatever challenges they faced — mites, weather, disease, nutrition.

By using these survivors as your breeder queens, you are conducting a selection program driven by real-world performance rather than theoretical ideals.

How to Develop a Survivor Stock Program

1. Stop treating your best colonies. This is the hardest step. Designate a portion of your apiary (20–30% of colonies) as your "selection yard." Do not treat these colonies for Varroa. Monitor their mite counts monthly.
2. Track survival through the full year. Colonies that survive a full annual cycle — including winter — without treatment are your candidates for breeding.
3. Evaluate secondary traits. Among the survivors, apply your scoring system for gentleness, honey production, and other traits. Not all survivors are worth propagating; some survived by luck or by being in a favorable location.
4. Rear queens from your top-scoring survivors. Use these queens to requeen your production colonies.
5. Repeat every year. Over time, the untreated portion of your apiary will be dominated by genuinely resistant stock.

Risks and Mitigations

The primary risk of a survivor stock program is colony loss. Losing 40–60% of your untreated colonies in the first few years is normal and expected. This is why the program should involve only a portion of your total apiary — your production colonies should continue to receive appropriate treatment and management.

The secondary risk is losing genetic diversity. If your survivor population drops to very few colonies, you may end up with a genetically narrow base that is vulnerable to inbreeding depression. Maintain at least 10–15 breeder colonies if possible, and periodically introduce outside genetics (from other resistant lines) to refresh your gene pool.

🌍 Perspective: Some of the most celebrated mite-resistant bee populations in the world — the feral bees of the Arnot Forest in New York, the Gotland bees in Sweden — became resistant through natural selection alone, with no human intervention. Survivor stock programs attempt to accelerate this process. Nature has already demonstrated that it works.

Evaluating Colonies for Breeding

The Evaluation Scorecard

A systematic scoring system transforms breeding from gut feeling into data-driven decision-making. Use the following scorecard at least twice per season — once during the main flow and once during late-summer or fall evaluation.

Calculating the Weighted Score

For each colony, multiply each trait score by its weight and sum the results:

Weighted Score = (Gentleness × 0.15) + (Honey × 0.20) + (Varroa Resistance × 0.25)
                 + (Winter Survival × 0.15) + (Brood Pattern × 0.10)
                 + (Spring Buildup × 0.10) + (Swarming × 0.05)

The maximum possible score is 5.0. Colonies scoring above 4.0 are strong breeder candidates. Colonies scoring below 2.5 should be considered for requeening.

What to Measure and When

Number of Evaluations Needed

Score each colony at least three times during the active season. A single evaluation is unreliable — a colony that scores well once may have had an unusually good day, or may be in a temporarily favorable forage position. Consistent scores across multiple evaluations are a much stronger indicator of genetic quality.

Culling: The Hard But Necessary Practice

Why Culling Matters

Selection is only effective if the bottom performers are removed from the breeding population. If you select your best queens but continue to allow your worst queens to produce drones, the genetic gains from your top colonies will be diluted by the inferior genetics contributed by the bottom.

Culling — removing or requeening colonies that do not meet your standards — is the necessary complement to positive selection.

What to Cull

Colonies that should be removed from the breeding population (either by requeening with better stock or by euthanizing the colony in severe cases):

1. Colonies with persistent high mite loads despite treatment — These are genetically susceptible and will produce susceptible drones.
2. Excessively defensive colonies — A colony that requires a full suit, heavy smoke, and still sends bees at your face is not worth keeping. The liability and enjoyment cost outweigh any production benefit.
3. Colonies that fail to build up in spring despite adequate resources — Chronic poor performers pull down your apiary average.
4. Colonies that swarm repeatedly despite proper management — High swarming tendency has a genetic component that you do not want to propagate.
5. Colonies with chronic disease (European Foulbrood, chalkbrood that does not resolve) — While some disease susceptibility is environmental, colonies that are persistently sick are poor breeding candidates.

Drone Culling

Drone culling is as important as queen culling. Because a single drone-producing colony can send thousands of drones to local DCAs, one below-average colony can contribute a disproportionate share of the genetics in your mating area.

Practical drone culling:

• Use drone foundation in your top colonies and worker foundation in the rest.
• Cut out drone comb from colonies scoring below your culling threshold.
• In extreme cases, move underperforming colonies to a separate location away from your mating yards during queen-rearing season.

Emotional Discipline

Culling is emotionally difficult. You invested time, money, and care in every colony. But livestock improvement requires removing the bottom performers to make room for the top. Every colony you keep that is below standard is a colony that produces drones that will mate with queens from your best stock — pulling the genetic average of your entire apiary downward.

Maintaining Genetic Diversity in Small Apiaries

The Inbreeding Problem

Honey bees are particularly vulnerable to inbreeding because of the complementary sex determination (CSD) locus. At this locus, heterozygous individuals develop as normal females (workers or queens). Hemizygous individuals (drones, with their single allele) develop as normal males. But homozygous individuals — those with two identical alleles at the CSD locus — develop as "diploid drones," which are non-viable and are consumed by workers at the larval stage.

In a small, closed breeding population, the number of distinct CSD alleles decreases over time through genetic drift. When a queen mates with drones carrying the same CSD allele she carries, 50% of her diploid offspring are non-viable diploid drones. This shows up as a "shot brood" pattern — a spotty, irregular laying pattern with many empty cells — and indicates serious inbreeding.

Strategies for Maintaining Diversity

1. Maintain at least 30 breeder colonies. Below this number, genetic drift causes rapid loss of CSD alleles. If your apiary is smaller than 30 colonies, you are breeding from a population that is too small to sustain long-term genetic health.
2. Introduce outside genetics every 2–3 years. Bring in queens or nucs from unrelated stock — ideally from a different breeding program or geographic area. This injects fresh CSD alleles and other genetic diversity.
3. Use multiple breeder queens each generation. Never breed exclusively from a single queen, no matter how outstanding she is. A single-queen breeding program is an inbreeding program by definition.
4. Rotate drone-producing colonies. Do not use the same colonies as drone sources year after year. Rotate your drone producers to ensure broad genetic representation.
5. Monitor brood pattern quality. Spotty brood that cannot be explained by disease, poor nutrition, or queen failure may indicate inbreeding. Take it seriously.

The 50/500 Rule

In conservation genetics, the "50/500 rule" suggests that a minimum effective population size of 50 is needed to prevent inbreeding depression in the short term, and 500 is needed for long-term genetic viability. For honey bees, where a single colony produces one queen but many drones, the effective population size is heavily influenced by the number of breeding queens.

For a small apiary (10–30 colonies), the practical implication is clear: you must bring in outside genetics regularly. You cannot run a closed breeding program at this scale without running into inbreeding problems within 5–8 generations.

Record-Keeping for Breeding Programs

Why Records Are Non-Negotiable

Breeding without records is guesswork. You cannot evaluate genetic trends, identify your best stock, or avoid inbreeding without systematic documentation. The difference between a beekeeper who "breeds" and a beekeeper who breeds is the quality of their records.

Minimum Required Data

For every colony, every season, record:

Tools for Record-Keeping

• Digital apps — Apps like Beetight, HiveTracks, and the CosmoBee platform allow structured data entry, automatic scoring calculations, and trend analysis across seasons.
• Spreadsheets — A well-designed spreadsheet is simple, portable, and fully customizable. Create one tab per season, one row per colony.
• Physical notebooks — Low-tech but reliable. Best used as a backup to digital records, not a replacement. Weather and ink do not mix well in the bee yard.

Whatever system you choose, consistency matters more than sophistication. Three years of complete data in a basic spreadsheet is far more valuable than one month of data in an elaborate app that you abandon.

Annual Breeding Report

At the end of each season, produce a brief breeding report:

1. Number of colonies evaluated and number meeting breeder criteria.
2. Average composite score for the apiary compared to previous years.
3. Top 3–5 breeder queens with their individual trait scores.
4. Colonies culled and reasons for culling.
5. Outside genetics introduced (source, number, integration results).
6. Plans for next season — trait priorities, target number of queens to rear, any stock introductions planned.

This report becomes your institutional memory. Without it, every season starts from scratch.

Resources for Bee Breeders

Books

• "Breeding Queen Bees" by John E. Eckert and Harry R. Shaw — A foundational text covering the biology and techniques of queen breeding.
• "Queen Rearing Essentials" by Lawrence John Connor — Practical, accessible guide focused on queen rearing methodology.
• "Honey Bee Biology and Beekeeping" by Dewey M. Caron and Lawrence John Connor — Comprehensive reference with extensive coverage of bee genetics.
• "The Beekeeper's Handbook" by Diana Sammataro and Alphonse Avitabile — Contains clear sections on queen rearing and stock selection.

Research Institutions

• USDA Honey Bee Breeding, Genetics, and Physiology Laboratory (Baton Rouge, LA) — The leading US institution for honey bee genetics research. Source of VSH stock and extensive publications.
• University of Minnesota Bee Lab — Pioneers of the Minnesota Hygienic line; excellent extension resources for beekeepers.
• University of Georgia Honey Bee Program — Active research on queen quality, breeding, and stock evaluation.
• Beelab at Penn State University — Research on honey bee health, genomics, and breeding.

Organizations

• American Beekeeping Federation (ABF) — Annual conference includes breeding workshops and presentations.
• Apiary Inspectors of America (AIA) — Resources on bee health and regulatory aspects of breeding.
• International Bee Research Association (IBRA) — Publishes the Journal of Apicultural Research, a primary venue for breeding research.
• State and local beekeeping associations — Many offer queen-rearing workshops and breeding-focused educational events.

Online Resources

• BeeSource.com — Active forums with dedicated sections on breeding and queen rearing.
• The Honey Bee Health Coalition — Toolboxes and guides for Varroa management, including breeding-based approaches.
• Managed Pollinator CAP (Coordinated Agricultural Project) — Research summaries and extension materials on honey bee health and genetics.

Practical Programs

• Bee Informed Partnership (BIP) — Runs the Tech-Transfer Team program, which provides breeding consultation and stock evaluation services to commercial beekeepers.
• State queen-rearing cooperatives — Several states have cooperative programs where beekeepers share breeder stock and coordinate mating areas. Check with your state apiarist or extension service.

References

1. Harbo, J. R., & Harris, J. W. (2005). Suppressed mite reproduction explained by the behaviour of adult bees. Journal of Apicultural Research, 44(1), 21–23.
2. Rinderer, T. E., et al. (2001). Resistance to the parasitic mite Varroa destructor in honey bees from far-eastern Russia. Apidologie, 32(4), 381–394.
3. Spivak, M., & Reuter, G. S. (2001). Resistance to American foulbrood disease by honey bee colonies Apis mellifera bred for hygienic behavior. Apidologie, 32(6), 555–565.
4. Seeley, T. D. (2007). Honey bees of the Arnot Forest: A population of feral colonies persisting with Varroa destructor in the northeastern United States. Apidologie, 38(1), 19–29.
5. Wallberg, A., et al. (2014). A worldwide survey of genome sequence variation provides insight into the evolutionary history of the honeybee Apis mellifera. Nature Genetics, 46(10), 1081–1088.
6. Page, R. E., & Peng, C. Y. S. (2001). Aging and development in social insects with emphasis on the honey bee, Apis mellifera L. Experimental Gerontology, 36(4–6), 695–711.
7. Delaney, D. A., et al. (2011). The physical, insemination, and reproductive quality of honey bee queens (Apis mellifera L.). Apidologie, 42(1), 1–12.
8. Oxley, P. R., & Oldroyd, B. P. (2010). The genetic architecture of honeybee breeding. Advances in Insect Physiology, 39, 83–118.
9. Büchler, R., et al. (2014). The influence of genetic origin and its interaction with environmental effects on the survival of Apis mellifera L. colonies in Europe. Journal of Apicultural Research, 53(2), 1–9.
10. Zayed, A., & Packer, L. (2005). Complementary sex determination substantially increases extinction proneness of haplodiploid populations. Proceedings of the National Academy of Sciences, 102(30), 10742–10746.