Exploring The Future of Varroa-Resistant Bee Genetics

Uncover the trends shaping The future of varroa-resistant bee genetics. Get insights into the latest research and advancements in varroa-resistant bee genetics.

Honey production and pollination services in the United States face a clear threat from varroa and its role as a DWV amplifier. This parasitic mite teams with deformed wing virus to create what many call a population-level “Monster” that drives colony losses.

Commercial practices can worsen that risk. Dense apiaries and yearly replacement with susceptible stock let mites drift and let virulent varroa–DWV pairings spread fast. Such cycles erode productivity for large operations and small hobbyists alike.

Long-term relief depends on shifting stock toward natural resistance. Practical breeding in apis mellifera aims to lower mite reproductive success inside brood rather than relying solely on miticides. That path pairs field selection with genomic tools to scale traits that stabilize colonies.

Key Takeaways

  • Varroa acts as a vector and amplifier for DWV, driving major losses.
  • Current practices can select for more virulent varroa–DWV dynamics.
  • Breeding to cut mite fecundity offers durable resistance.
  • Apis mellifera selection combines field tests with genomic metrics.
  • Scaling resistant stock reduces chemical dependence and boosts honey yields.
  • Report previews include mechanisms, GWAS, and a U.S. roadmap.

Why Varroa Still Matters: The Evolving Varroa/DWV “Monster” in U.S. Beekeeping

A shifting partnership between mites and DWV now dictates colony survival. That pairing shortens lifespan and cuts honey yields even when late treatments are applied.

From parasitic mite varroa to DWV synergy: what changed

Varroa acts as both vector and amplifier for DWV. When mites invade brood, viral loads climb and pupae emerge deformed or fail entirely. This interaction now drives many winter losses for honey bee operations.

Commercial apiary practices that select for virulence

Dense, multi-hive yards and rapid turnover of stock create highways for the most virulent mite–virus combos. Collapsing colonies shed virus-laden mites that reinvade neighbors. Annual replacement with susceptible queens keeps selecting for higher infestation rates.

  • 1987 arrival of varroa jacobsoni (now varroa destructor) spread quickly and led to early pyrethroid resistance and comb residues.
  • Biannual treatments remain common, yet losses persist—showing a need for true resistance, not only control.

Way forward: select traits that cut mite reproduction inside brood and pair assays with regional testing to limit reinvasion and stabilize populations.

Lessons from Apis cerana: A Template for Coexistence with Varroa destructor

Apis cerana offers clear, compact strategies that keep varroa and mite loads low while supporting healthy colonies.

Drone-focused reproduction and worker defenses

In cerana, mites reproduce mainly in drone brood. Worker cells resist mite feeding and cut mite reproduction sharply.

Social apoptosis and early removal

Social apoptosis occurs when worker larvae detect mite salivary cues and trigger self-removal. Nurse bees then remove infested brood before mites can complete a cycle.

Sniffing holes and entombment

Workers keep a small inspection hole in drone cappings to monitor pupal health. When pupae are stressed, bees may entomb the cell, trapping mites in a reproductive dead end.

  • Result: mites face strong costs when reproduction is limited to drones.
  • Evolutionary logic: this aligns parasite fitness with host survival and enforces prudence in mites.
  • Applied goal: breed apis mellifera that detect and remove infested worker brood, while managing drone cycles to contain mite spread.

The future of varroa-resistant bee genetics

Breeding programs now aim to steer natural selection toward traits that curb mite reproduction inside worker brood.

Strategic pivot: shift effort from repeated chemical knockdowns to selecting honey bees that lower varroa daughters per foundress. That change reduces population growth of destructor without relying on constant treatments.

Natural selection and parallel evolution

Resistant populations in South Africa and South America show parallel paths—recapping, removal, and suppressed mite reproduction. These independent cases suggest repeatable traits that breeders can target.

Why reduce reproduction, not only tolerate mites

Tolerance can let mite loads climb, which raises DWV risk and can trigger colony collapse. Small fecundity shifts—halving average daughters per cycle—drive mite numbers below replacement in simulation models.

  • Select traits that lower mite reproduction in brood.
  • Prioritize heritable colony-level behaviors that maintain honey production and stable colonies.
  • Design U.S. selection programs around regional testing, measured traits, and realistic mating control.

Core Resistance Mechanisms: Hygienic Behavior, VSH, Recapping, and SMR/DMR/MNR

A handful of social defenses explain why some colonies keep mite loads low.

Varroa sensitive hygiene (VSH) means worker bees detect and remove infested brood cells before mites finish a reproductive cycle. Colonies with ~50% VSH can hold varroa growth in check in simulation and field tests. This trait targets reproductive cells and drops mite reproduction quickly.

A detailed and dynamic illustration showcasing hygienic behavior in honeybees. In the foreground, a close-up view of a worker bee meticulously grooming and cleaning another bee, reflecting the intricate process of VSH (Varroa Sensitive Hygiene). In the middle ground, a group of bees collectively tending to the hive, with some removing debris and others inspecting the comb. The background depicts the interior of the hive, with a warm, amber-toned lighting that creates a sense of activity and industry. The composition emphasizes the cooperative nature of the hive's hygiene and the critical role it plays in maintaining bee health and resistance to Varroa mites. Rendered in a detailed, naturalistic style to capture the nuances of this vital behavior.

Recapping and targeted uncapping

Recapping is a low-cost social immune action. Workers open and then re-seal cappings, disrupting mite egg laying without killing pupae. Experimental uncapping shows causal drops in mite fertility and it scales well in selection programs.

Suppressed mite reproduction (SMR/DMR/MNR)

SMR/DMR/MNR describe brood-level effects that reduce mite fertility inside cells. Survivor stocks historically show these patterns, and they cut mean daughters per foundress—an Achilles heel for mite population growth.

Grooming and biting

Grooming and mandibular biting help remove phoretic mites, but studies find they rarely suffice alone in managed apiaries. Use grooming as a supportive trait while you stack VSH, recapping, and MNR in selection to depress mite reproduction across seasons.

  • Screening: freeze or pin-kill tests for hygienic behavior, measure recapping hit rates, inspect cocoons for entombment, and track MNR.
  • Limits: grooming aids control varroa but is inconsistent as a standalone defense.
  • Strategy: stack complementary traits in apis mellifera to build durable resistance and stabilize honey production.

New Evidence Shaping 2025: Melissa Oddie’s Findings on Recapping and Mite Fertility

Recent field work by Melissa Oddie links high recapping rates with sharp drops in mite fertility across several populations. This research focuses on worker behavior that interrupts reproduction inside worker cells and measures colony-level effects.

Metrics and population patterns

Resistant colonies show elevated recapping (~77%) and brood removal (~80%) of infested brood. Across sites, those behaviors correlate with low mite fertility (r ≈ 0.77).

Metric Value Biological effect
Recapping rate ≈77% Disrupts mite egg laying in worker cells
Brood removal ≈80% Lowers surviving offspring per foundress
Correlation (recap vs fertility) r ≈ 0.77 Strong negative association across populations

Proven causality and operational value

Experimental uncapping isolated recapping’s effect and showed causality: recapped cells produced fewer mite offspring without killing nestmates. This makes recapping a low-cost social immunity action.

Why this matters: recapping preserves workforce and honey production while cutting mite population growth. Pairing recapping with vsh and MNR offers multi-mechanism suppression of mite reproduction in managed colonies.

Breeding takeaways: score lines for recapping during peak brood, compare offspring counts per foundress in recapped versus untouched cells, and prioritize high-recapping apis mellifera stock for propagation.

Genetics at Scale: 2024 GWAS Shows Resistance Is Heritable and Polygenic

A continent-wide genome scan of more than 1,500 apis mellifera colonies used pool sequencing and inferred queen genotypes to link colony phenotypes to DNA.

Study design: researchers combined whole-genome pool-seq of workers with queen genotype reconstruction. They scored three colony-level traits: recapping, mite non reproduction (MNR/DMR), and overall infestation.

Key finding: sixty significant associations appeared, but no large-effect locus emerged. This shows resistance is substantially heritable and polygenic rather than a single-gene switch.

Why those traits matter

Recapping, MNR, and infestation measure how colonies suppress mite reproduction inside cells. Together they capture social and brood-level defenses that keep populations low.

Element Data Implication
Sample size 1,500+ colonies High power to detect small-effect loci
Phenotypes Recapping, MNR, infestation Colony-level biology, not single bees
Genetic architecture 60 associations, polygenic Genomic selection preferred

Practical takeaway: genomic prediction can cut costly field scoring, flag promising queens early, and guide crosses to avoid inbreeding. With tailored markers and meta-analyses, regional selection programs can scale resistant stock for honey producers and hobbyists across populations.

Breeding for the Real World: Controlling Drone Pools, Isolation, and Instrumental Insemination

Practical breeding succeeds when mating is managed, not left to chance. Queen and package producers must adopt clear mating plans to protect gains in resistance.

Producer responsibilities in the U.S. market

Producers should commit to isolated mating yards, targeted drone flooding from proven lines, or instrumental insemination for key queens. These steps reduce open-mating dilution and help seed regions with hardy stock.

Assays that matter

Prioritize simple, repeatable tests: freeze or pin-kill assays for hygienic response, recapping rate checks, quick cocoon inspections for entombment, and mite fertility or MNR counts per foundress. Track results by queen line and mating record.

Action Why it matters Practical target
Isolated mating yards Limits unwanted drone gene flow Maintain 3–5 km buffer or use island sites
Instrumental insemination Ensures matched drones from resistant mothers Use for nucleus and breeder queens
Routine assays Verify colony-level resistance Quarterly freeze/pin-kill, MNR sampling

Scaling and logistics: form regional cooperatives, time drone rearing with queen flights, and publish mating and phenotype records. Customers gain verifiable honey and colony stability, lower treatment costs, and access to robust honey bees suited to U.S. operations.

Population Dynamics: Why Resistance Is a Population Trait, Not Just a Colony Trait

A lone resistant colony can fail fast if surrounding populations flood it with mites.

Resistance scales. One hive that shows strong behavior can still lose ground when nearby colonies collapse and shed large mite loads. Reinvasion from heavy infestation drives rapid rises in varroa numbers and raises DWV risk for honey production and colony survival.

Mite drift and robbing matter. Foragers and robbers carry phoretic mites between yards. This movement undermines gains from careful breeding unless neighbors also express similar traits.

A detailed close-up view of a varroa mite population thriving on the surface of a honeycomb. The mites are shown in varying shades of red and brown, with intricate textural details and translucent bodies, clinging to the geometric hexagonal comb structure. The lighting is soft and natural, casting gentle shadows that accentuate the mites' forms. The background is slightly blurred, with the faint outlines of other comb cells visible, creating a sense of depth and context. The overall tone is one of scientific observation, allowing the viewer to study the complex dynamics of this parasitic mite species and its impact on honey bee populations.

Mite drift, reinvasion, and vulnerability

Resistant colonies can be overwhelmed when placed into mite-dense regions. Sudden influxes can defeat partial resistance, especially if brood removal rates already exceed safe bounds.

Designing apiaries to preserve gains

Practical layout cuts exchange pathways. Increase spacing, lower colony density, and reduce robbing cues like exposed syrup or weak hives.

  • Cluster resistant colonies together to create local buffers.
  • Use buffer zones with managed, low-infestation stock around core yards.
  • Time brood breaks across yards to lower peak mite drift seasons.

“Survivor populations depend on community-wide trait expression; isolated wins rarely hold without coordinated placement and monitoring.”

Risk factor Why it matters Practical countermeasure
Mite drift Moves mites between colonies and yards Increase spacing, reduce robbing cues
Reinvasion Overwhelms partial resistance Cluster resistant colonies; use buffer zones
High brood removal Can weaken isolated colonies during influx Coordinate brood breaks; monitor mite counts

Monitor and adapt. Track mite movement and adjust placement or interventions. Regular checks keep honey yields steady and help apis mellifera populations retain useful traits over time.

Global Case Studies: Cuba, South Africa, Brazil—Natural Selection in Action

Real-world populations offer clear proof that low-intervention programs can fix useful traits at scale.

Cuba maintains roughly 220,000 treatment-free colonies for more than twenty years. These populations produce about 40–70 kg honey per hive and are noted for mild temperament that eases management. Such scale shows that large numbers of apis mellifera can deliver stable honey yields without routine chemical controls.

In Brazil and South Africa, researchers link lower DWV loads to strong rates of brood removal. More than half of infested pupae are removed before emergence in some sites. That action cuts viral pressure on adult workers and lowers colony-level infestation.

Across these populations, elevated recapping and higher brood removal match reduced mite fertility. Parallel selection produced similar traits in different environments, which supports natural selection as an effective force for resistance.

  • Policy lesson: minimal interference lets traits spread fast across populations.
  • Management takeaway: prioritize regional selection and verified field performance when scaling resistant stock.
  • U.S. implication: population-wide programs, not lone-colony trials, give the best chance to protect honey production.

Feral Bees and Small Nests: Tom Seeley’s Arnot Forest Insights

Arnot Forest research shows how simple life-history traits sustain stable honey production without treatment.

Small cavities, frequent swarming, and low density reduce continuous brood and shorten windows for varroa reproduction. Smaller nest volumes mean fewer brood cycles at once. That limits mite breeding and slows population growth.

Established colonies lived about five to six years on average. Annual queen turnover via swarming creates regular brood breaks. Those breaks are natural pauses that cut mite offspring production.

What managers can copy

Practical steps include using smaller hive volumes, planned brood interruptions, and spacing that lowers colony density. These moves mimic feral dynamics and reduce reinvasion risk.

Ecology and organization matter alongside genetics. While selection for resistance helps, colony layout, swarming rhythm, and habitat shape long-term outcomes more than single genes in many cases.

Feature Effect on mite Management tip
Small nest volume Fewer simultaneous brood cells Use nucs or smaller boxes
Frequent swarming Regular brood breaks Allow controlled splits or timed brood removal
Low colony density Less mite drift and robbing Space yards; cluster resistant colonies

Managing the Transition: From Biannual Treatments to Selective, Minimal Intervention

Shifting away from blanket twice-yearly treatments is possible, but it must be managed without inviting large winter losses. In the U.K., most keepers treat—often two times a year—and long-term overwinter loss averages near 18%. A small minority remain treatment-free for six years or more, showing a plausible path forward.

Trends and realistic baselines

Most producers in temperate zones still rely on treatment to hold mite numbers and protect honey yields. In the U.S., survey data mirror that pattern—treatment-free operations are rare.

Practical steps to avoid a death spiral

Selective intervention means treating only outlier colonies with high mite counts to protect neighbors while keeping selection pressure. Use consistent sampling and record infestation rates.

Biomechanical controls—planned brood breaks, drone brood removal, and heat or temperature tactics—cut reproduction without chemicals. Sync actions with nearby keepers, reduce robbing, and time interventions before collapse season.

  • Catch persistent feral swarms and assess before adding to yards.
  • Coordinate treatment timing across local operations.
  • Track mite counts, recapping, and removal rates to guide decisions and selection.

For seasonal guidance and practical chores, consult a schedule of seasonal tasks to align interventions with honey production cycles and population-level goals.

Targeting Mite Reproduction: The Achilles’ Heel Inside Brood Cells

Modest tweaks to postcapping timing and hive temperature can compress the window varroa needs to lay eggs. Shortening that window lowers average mite reproduction and slows population growth.

Shortening postcapping duration and temperature tuning

Varroa fecundity is higher in drone cells because postcapping lasts ≈15 days versus ≈12 in worker cells. Compressing postcapping by a day or raising broodnest heat a few degrees can cut daughters per foundress.

Drone brood management

Limit drone brood timing and volume. Removing or timing drone frames reduces the most fecund pathway for mite reproduction and helps nearby colonies and honey bees stay productive.

Entombment under the cocoon: a practical trait to screen

Jeff Harris documented larvae trapping foundress mites beneath cocoon silk. Inspecting cocoons in low-mite colonies reveals trapped foundresses quickly.

Practical checklist:

  • Modulate postcapping timing during peak brood.
  • Schedule drone frame removal to cut mite fecund cycles.
  • Screen cocoons for entombed foundresses as a fast assay.
  • Combine entombment, recapping and VSH to push varroa below replacement.
Lever Mechanism Expected effect
Shorter postcapping Less time for egg laying Lower mite reproduction rate
Temperature tuning Raise broodnest by 1–2°C Reduced daughters per foundress
Drone brood control Limit and time frames Cut highest fecund reproduction pathway
Entombment screening Inspect cocoons for trapped mites Rapid selection marker for colonies

DWV Management Through Bee Genetics: Reducing Viral Burden Indirectly

Reducing virus loads in colonies requires tactics that cut mite-driven amplification while preserving brood and honey yield. Genetics alone won’t stop transmission, but colony-level behaviors can shift viral equilibria as mite pressure falls.

Colony-level behaviors that limit DWV despite mite presence

Recapping interrupts mite reproduction without destroying pupae. That lowers the number of new mites that carry high viral loads into adult workers.

Targeted brood removal in Brazil and South Africa links to lower DWV at the colony level when removals outpace mite reproduction. Removing infected pupae reduces nidus points for virus amplification.

Cannibalization and trophallaxis: managing trade-offs as VSH increases

Hygienic removal and VSH cut mite reproduction, but cannibalizing infected brood can temporarily raise DWV in nurse bees via trophallaxis.

Over time, as resistance traits lower mite numbers, that transient rise falls and a new, lower viral equilibrium emerges. Monitoring helps confirm that trend before scaling out stock.

Operational balance: favor recapping-heavy lines to disrupt mite reproduction while avoiding excessive brood loss from aggressive cannibalization. Combine recapping, measured hygienic behavior, and grooming to reduce both infestation and viral amplification.

Intervention Primary effect Risk / trade-off
High recapping rate Disrupts mite egg laying; preserves brood Low risk to honey yield
Targeted brood removal Removes high-DWV pupae Transient adult DWV uptick via trophallaxis
VSH / hygienic behavior Reduces mite reproduction Possible brood loss if overexpressed
Combined monitoring Tracks mite fertility and viral load Requires lab support for virus quantification

Monitoring recommendation: pair mite fertility counts with periodic DWV quantification to validate that selected traits lower colony viral burden over time. That evidence helps producers scale resistant stock while protecting honey production and colony health.

Roadmap for Genomics-Enabled Breeding in the United States

Genomic tools can speed selection when paired with standardized field assays and coordinated mating yards. This roadmap links lab pipelines to practical steps that producers can adopt across regions.

From pool-seq to marker panels

Start with pool-seq of worker samples and queen genotype reconstruction to run colony-level GWAS for recapping, MNR, and infestation.

Early wins come from marker-assisted selection using high-impact panels while building larger training sets for genomic prediction.

Regional mating and tester networks

Operationalize selection by creating drone-flooded yards and calibrated tester apiaries. Flooding reduces open-mating dilution and preserves selected traits in local populations.

  • Standardize phenotyping: consistent protocols for recapping, VSH, MNR, and infestation across tester sites.
  • Build feedback loops: feed field performance into genomic models to prioritize lines that balance resistance and honey productivity.
  • Scale: shift from marker panels to genomic predictions as datasets exceed thousands of colonies.

“Integrating pool-seq, queen inference, and regionally controlled mating turns associations into usable selection tools.”

For program design and federal research links, consult using genetics to improve breeding and health of honey.

Market, Policy, and Timeline: What Could Accelerate Resistant Stock Adoption

Certification programs and clear performance data create market trust and speed adoption of resistant lines. Buyers who want lower treatment bills and steady honey yields will pay for queens that carry verified resistance traits.

Consumer demand and verification

Verified performance matters. Independent regional testing networks can publish standard results for recapping, MNR, VSH, and infestation outcomes.

Drivers include:

  • Queen buyers seeking lower cost and better overwintering for colonies.
  • Retail and packers preferring honey from verified, low-treatment operations.
  • Volunteer testing panels that function like an “All-America” program to validate claims.

For supply-chain analysis and sector planning, see a detailed queen sector analysis.

Three scenarios to 2030

Scenario one: slow, incremental gains as selection spreads through committed breeders and regional mating yards.

Scenario two: a tipping point where major queen buyers demand certified lines and adoption accelerates nationwide.

Scenario three: regionalized resistance where local apis mellifera populations adapt separately and trade in resilient queens remains local.

“Market pull, backed by verification and policy support, can flip selection pressure across an entire population.”

Policy levers such as grants for breeder infrastructure, support for isolated mating yards, and incentives for data sharing will shorten timelines and help producers scale resistant bees while protecting honey production.

Conclusion

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Stacking social and brood-level traits offers a realistic route to stable, productive colonies. Practical work shows that lowering mite reproduction across brood cycles cuts varroa growth while protecting honey and worker numbers.

Evidence converges: natural populations, experimental uncapping, and large-scale GWAS point to polygenic social immunity as a durable path to resistance in apis mellifera.

Execution needs coordinated action: breeders must align selection with regional drone control, repeatable assays, and genomic tools. Build tester networks and verification programs so buyers can choose proven stock.

Act now to scale verified lines, protect honey yields, and help U.S. bees recover across local populations.

FAQ

What is varroa destructor and why does it matter for Apis mellifera?

Varroa destructor is a parasitic mite that infests honey bee (Apis mellifera) colonies. It weakens individual bees by feeding on hemolymph and spreads viruses such as deformed wing virus (DWV). High mite loads cause colony decline and large-scale losses in commercial and hobby apiaries. Managing mite reproduction and reinvasion is central to colony health.

How do hygienic behaviors like VSH and recapping reduce mite reproduction?

Varroa sensitive hygiene (VSH) targets and removes infested brood cells, directly interrupting mite reproduction cycles. Recapping involves workers opening and resealing brood cells, which disturbs mite mating and lowers fertility without killing pupae. Both behaviors lower mite population growth and reduce viral transmission at the colony level.

Are grooming and biting effective resistance traits on their own?

Grooming and biting support colony defense by removing some phoretic mites from adult bees, but they rarely stop infestations alone. When combined with brood-based traits such as VSH, recapping, and suppressed mite reproduction (SMR), grooming increases the overall resilience of the colony.

What is suppressed mite reproduction (SMR/DMR/MNR) and how is it measured?

Suppressed mite reproduction refers to reduced mite fertility inside brood cells. Researchers score the proportion of foundress mites producing viable offspring. Lower mite fertility indicates SMR/DMR/MNR and predicts slower population growth. Standard assays count mite offspring per cell and record the presence of mature female progeny.

How heritable are resistance traits and what does polygenic mean for breeders?

Recent genome-wide association studies (GWAS) show resistance traits are heritable and polygenic—driven by many loci with small effects. That makes genomic selection and multi-trait phenotyping more effective than selecting single genes. Breeders should use marker-assisted and genomic tools alongside phenotypic assays.

Can beekeepers rely on natural selection alone to produce resistant stock?

Natural selection can yield resistant populations, as seen in Cuba, Brazil, and South Africa. However, achieving reliable, widespread resistance takes time and favorable population dynamics. Active breeding, controlled mating, and apiary design speed progress and reduce reinvasion risks.

Why is resistance a population trait rather than just a colony trait?

Mite drift, robbing, and drone-mediated gene flow cause reinvasion between colonies. A single resistant colony surrounded by susceptible neighbors will face persistent mite pressure. Maintaining resistance requires coordinated regional efforts, reduced local mite loads, and apiary layouts that limit drift.

What practical breeding measures reduce inadvertent selection for virulence?

Control drone pools, use isolated mating yards or instrumental insemination, and adopt multi-phenotype selection (recapping, VSH, MNR, infestation metrics). Verify queens and packages for resistance traits and avoid importing high-mite stock that can push selection toward greater virulence.

How does targeting mite reproduction compare to killing mites with treatments?

Targeting reproduction lowers long-term population growth and viral amplification without heavy chemical reliance. Treatments reduce mites quickly but can select for resistance in mites and harm bees. Integrating selective treatment with breeding for reduced mite fertility offers a sustainable path.

What role does recapping play according to recent research?

Studies, including Melissa Oddie’s work, link high recapping rates to reduced mite fertility across locales. Experimental uncapping shows causality: disturbing brood cells lowers mite reproductive success while preserving pupae, making recapping a low-cost social immunity trait to select for.

How should commercial queen producers adapt to support resistant stock?

Producers must test for hygiene, recapping, and mite fertility, manage drone sources, and adopt genomic selection tools. Transparent verification and consistent assays help buyers choose queens that perform under real-world conditions. Package and queen producers also need to limit mite loads before sale.

Can genetic approaches reduce DWV impacts even if mites persist?

Yes. Colony-level behaviors that remove or limit infested pupae reduce viral replication and transmission. By lowering mite fertility and brood infestation, breeding can reduce DWV prevalence and severity indirectly, improving colony survival despite some mite presence.

What apiary management steps help preserve resistance after breeding?

Design apiaries to limit drift, regularly monitor mite levels, use selective rather than blanket treatments, and coordinate with neighboring beekeepers. Maintaining small cavity sizes, encouraging swarming behavior when feasible, and protecting feral-type nesting dynamics can also support resistance.

Are there real-world examples where treatment-free beekeeping succeeded?

Yes. Cuba reports large treatment-free apiaries with high honey yields after natural selection favored resistant traits. Brazil and South Africa document lower DWV loads tied to increased brood removal. These cases show selection can work at scale when reinvasion is limited and selection pressures persist.
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