Bees hanging from each other in a festooning chain

Honey bee genetics: inheritance, heritability, controlled mating and breeding for varroa resistance in the UK

Honey bee genetics for UK beekeepers: the csd gene, multiple mating, heritability, hygienic and VSH bees, the native black bee and how to improve your stock.

Honey bee genetics rests on three facts that make bees unlike any farm animal: drones are haploid and hatch from unfertilised eggs, sex is decided by a single gene called csd, and a queen mates with a dozen or more drones on a few flights early in life. The traits you care about, honey yield, temper, swarming and resistance to varroa, belong to a whole colony with one mother and many fathers. That is why improving bees needs controlled mating, records on many related colonies and patience, and why “varroa resistant” queens so often disappoint once they are open mated in a British apiary.

This guide explains how inheritance works in the hive, how heritable the main traits are, what the best breeding programmes have achieved, how mating can be controlled in the UK, what genetics says about the native dark bee, and what a hobby beekeeper with five or ten National hives can realistically do.

How does honey bee genetics work?

Haplodiploidy: drones without fathers

The western honey bee, Apis mellifera, has 16 pairs of chromosomes in females (2n = 32) and a single set in males (n = 16). Each time the queen lays, she decides whether to release sperm from her spermatheca onto the egg. The University of Florida’s IFAS profile of the European honey bee summarises the result:

  • Fertilised egg: diploid, it becomes a female. Larval feeding then decides whether she develops as a worker or a queen, as our page on the queen bee explains.
  • Unfertilised egg: haploid, it becomes a drone, which carries only its mother’s genes.

The same IFAS profile gives the development times that every queen rearer works to. All three castes spend 3 days as an egg. The larval stage then lasts about 5.5 days for a queen, 6 for a worker and 6.5 for a drone, and the pupal stage about 8, 12 and 14.5 days. Egg to adult takes 15 to 16 days for a queen, 21 for a worker and 24 for a drone. Our page on the honey bee life cycle follows each stage in detail.

For a breeder, the key consequence is that a drone passes its whole genome, unshuffled, to every daughter. All sperm from one drone are genetically identical.

The csd gene and diploid drones

In 2003 Martin Beye, then at Martin Luther University Halle-Wittenberg in Germany, and Robert Page’s team at the University of California, Davis, reported in Cell the gene that sets sex in honey bees: csd, the complementary sex determiner. The mechanism works like this:

  1. A female carries two different csd alleles, one from each parent. Their two proteins combine into an active form that switches on female development.
  2. An unfertilised egg has only one copy of csd, so no active form appears and the embryo becomes male.
  3. A fertilised egg that receives two identical copies of the same allele behaves like an unfertilised one. It becomes a diploid drone, which is sterile, and workers detect and kill it as a young larva.

The original release mentioned 19 alleles. A later study led by Martin Hasselmann at the University of Cologne, published in Molecular Biology and Evolution in 2013 with S. Lechner as first author, found at least 87 alleles worldwide, 53 of them in Kenya alone, and estimated that the species holds 116 to 145 in total. That diversity sets a hard limit on inbreeding.

What diploid drones cost a colony

Suppose a queen carries alleles A1 and A2. If one of the drones she mated with carries A1, half the fertilised eggs from that drone’s sperm will be A1/A1 and will be removed. With a dozen fathers contributing roughly equally, one “matched” drone costs a few per cent of the worker brood. A queen inseminated with a single matched drone loses half her female brood. The result is a scattered, patchy brood pattern and a colony that never builds up.

It is a warning sign of a narrow population: the fewer csd alleles in circulation, the more likely a match. A patchy pattern can also come from an old queen or from brood disease, so rule those out first. American and European foulbrood are notifiable under the Bee Diseases and Pests Control Orders: report suspicion to the National Bee Unit (NBU) or your local bee inspector in England and Wales, to the Scottish Government’s Bee Inspectors in Scotland ([email protected]), with samples analysed by SASA, or to DAERA in Northern Ireland.

Why does the queen’s multiple mating matter?

Subfamilies, super-sisters and half-sisters

A virgin queen makes one or a few mating flights to drone congregation areas in her first two weeks, and, according to the IFAS profile, mates with about 15 drones. The sperm is stored in her spermatheca for life. A colony is therefore a patchwork of subfamilies, each made of workers sharing one father.

Relationship between two workers Coefficient of relatedness Why
Same father (super-sisters) 0.75 they share the whole paternal genome and half the maternal one
Different fathers (half-sisters) 0.25 only the maternal half is shared
Worker and her mother 0.5 as in any diploid animal

Multiple mating is not a curiosity. Heather Mattila and Thomas Seeley at Cornell tested it in Science in 2007, comparing 12 queens each inseminated with sperm from 15 drones against 9 queens each inseminated with one drone. According to Cornell’s summary, after two weeks the genetically diverse colonies had built about 30% more comb, stored 39% more food and foraged 27% to 78% more. By the end of summer they had about five times as many bees and eight times as many drones. A year after founding, about 25% of the diverse colonies were alive. Every uniform colony had starved.

Queen effects and worker effects

To a geneticist, every colony trait has two parts:

  • The queen effect (maternal effect): her laying rate, her pheromones, her own genes.
  • The worker effect (direct effect): the collective behaviour of daughters who each have a different father.

Modern breeding models estimate these separately. For the beekeeper the lesson is simple: you never evaluate a queen alone, you evaluate a queen plus the drones she mated with, and her daughters’ colonies decide whether she was good.

How heritable are honey bee traits?

Narrow-sense heritability (h²) measures the share of differences between colonies that is explained by genes passed on additively. The higher it is, the more a colony’s own performance tells you about its offspring. It describes a population, measured in a given environment by a given method. It says nothing about whether a particular line is good.

Published estimates

Trait Estimated heritability Population and source
Honey yield 0.02 (colony effect) to 0.11 Swiss dark bee and Carnica, Guichard et al., Apidologie, 2020
Calmness on the comb 0.12 to 0.16 Swiss dark bee, Guichard et al., 2020
Defensive behaviour 0.32 to 0.34 Swiss dark bee, Guichard et al., 2020
Swarming drive 0.06 to 0.07 Swiss dark bee, Guichard et al., 2020
Hygienic behaviour (pin test) 0.18 to 0.19 (dark bee); 0.06 to 0.09 (Carnica) Guichard et al., 2020
Varroa infestation and mite growth close to zero Guichard et al., 2020
Suppression of mite reproduction 0.46 Harbo and Harris, Journal of Economic Entomology, 1999

The Swiss figures are especially useful to British readers because one of the two populations is the dark European bee, Apis mellifera mellifera, the same subspecies as the native British black bee.

How to read these figures

  • Estimates are uncertain. The Swiss authors studied about 1,000 colonies in each population, with records from 2009 to 2018, and point out that standard errors were often as large as the estimates themselves. Treat every value as a rough guide.
  • Mite counts carry almost no genetic signal. In both Swiss populations, varroa infestation showed practically no additive genetic component. Drifting, robbing, brood levels and sampling swamp it. A colony with a low mite count in August may simply have had fewer neighbours.
  • Narrowly defined traits are better inherited. John Harbo and Jeffrey Harris obtained 0.46 for suppression of mite reproduction from 28 colonies in 7 groups of 4, each group headed by queens inseminated with semen from the same sire colony. They measured a precise mechanism inside brood cells, not an overall mite count. That reasoning led to VSH bees.
  • Temper responds well. Defensive behaviour is one of the more heritable traits, which is why selecting against bad temper works in a few generations when mating is controlled.

What can a breeding programme achieve? Lessons from BeeBreed

Breeding values and BLUP

Large programmes rank queens by breeding values calculated with BLUP (best linear unbiased prediction), the method developed by Charles Henderson for dairy cattle. BLUP combines a colony’s own performance with that of all its relatives and corrects for apiary and year, while estimating queen and worker effects separately.

Twenty-five years of Carnica data

The best documented example is BeeBreed, the European database for Apis mellifera carnica run from the Länderinstitut für Bienenkunde at Hohen Neuendorf in Germany. Andreas Hoppe and colleagues reviewed it in Insects in 2020:

Indicator Value
Queens with breeding values (14 February 2020) 224,081, of which 195,458 were performance tested, in 12 countries
Model BLUP, queen and worker effects; four-trait model (honey, gentleness, calmness, swarming) and two-trait varroa model
Index scale mean 100, standard deviation 10, based on queens tested in the last five years
Varroa index hygienic behaviour and varroa infestation development combined 57:43
Gain in honey yield, 1992 to 2018 more than 5 kg
Yearly gain, 2009 to 2018 about 336 g of honey; 0.73 percentage points of pin test clearance
Average inbreeding peaked at up to 10% around 1980, about 4% since 2000

Why it worked. Common test protocols, controlled mating at mating stations, a central register of queens and breeders that made exchange easy, and inbreeding monitoring all mattered alongside the statistics.

Its limits, stated by the authors. The correlation between breeding values and later performance was modest: 0.15 for honey yield, about 0.25 for gentleness and calmness and 0.17 for swarming. Hygienic behaviour was the best predicted trait, and varroa infestation development the worst. Varroa traits were recorded in only about 15% of all performance tests.

There is no equivalent national breeding database in the UK. British selection rests on individual breeders, bee farmers and voluntary groups.

How do you control mating in the UK?

Without control of the drones, half the genes in your workers are chosen by the neighbourhood. Three approaches exist.

Method Control of paternity Strengths Limits
Open mating none free, well mated queens local drones dilute any selected trait in one generation
Isolated mating site good to very good, depending on isolation many drones, sister drone colonies few truly isolated sites in Britain
Instrumental insemination complete planned crosses, single drone possible skill, cost, inbreeding risk in a closed population

How far drones travel

The best British data come from the Peak District. Jensen and colleagues used DNA paternity tests in 2005 in Edale and Hope Valley, two valleys chosen for dark bee conservation. Half of matings took place within 2.5 km of the mating apiary and 90% within 7.5 km, and one queen mated about 15 km away. The Losehill ridge between the valleys did not isolate them completely, although Edale stayed fairly isolated: in its most isolated apiary about 60% of matings involved Edale drones. In lowland England, with a hive in every other garden, true isolation is close to impossible.

Isolated sites and Colonsay

Islands, remote glens and open moorland without other colonies are the safest mating sites. The history is long: Hoppe and colleagues date the first known mating station to 1870, at Hroby in today’s Czech Republic, and the oldest still working, Gehlberg in Germany, to 1911. After multiple mating was discovered in 1954, stations came to be stocked with drone colonies headed by sister queens, so that the father side is at least a known line. Brother Adam’s Dartmoor mating apiary for the Buckfast bee worked on the same principle.

In Scotland, one such site is protected by law. The Bee Keeping (Colonsay and Oronsay) Order 2013, in force since 1 January 2014, prohibits keeping any honey bee other than Apis mellifera mellifera on the two islands, which makes them both a reserve for the native bee and a natural mating station.

Drone flooding

Where isolation is impossible, the practical tool is drone flooding: put drone comb into your chosen colonies, remove it from poor ones, and if possible requeen nearby colonies with your better stock. BIBBA’s bee improvement groups rely on this, and members agree not to import bees or queens or to propagate from recently imported stock. It tips the odds rather than guaranteeing the father.

Instrumental insemination

Insemination lets you choose the father exactly, down to a single drone. Single-drone crosses are used to build uniform lines or to test a particular sire, as in the VSH programmes. The cost is skill, equipment and, in a closed population, faster inbreeding: one father per queen reduces the number of csd alleles carried forward much more quickly than natural multiple mating. Our guide to instrumental insemination covers the method.

Can you breed varroa resistant bees?

Three traits not to confuse

  • Hygienic behaviour: workers detect, uncap and remove dead or diseased brood. It protects against chalkbrood and American foulbrood, and to a degree against varroa. The NBU described its freeze-killed brood test in Bee Craft in 2002: freeze a section of sealed worker brood for 24 hours, return it to the brood nest and count after 48 hours. A colony counts as hygienic if it removes more than 95% of the dead brood. In the NBU’s tests, 6 of 37 colonies (16%) qualified in 2000 and 2 of 41 (5%) in 2001.
  • Varroa sensitive hygiene (VSH): a targeted form of hygiene aimed at cells where mites are reproducing. According to eXtension, the US extension network’s bee health resource, workers about 15 to 18 days old detect infested brood and uncap it, and other bees then remove the pupa. The mite’s offspring die; the mother mite often survives.
  • Suppression of mite reproduction (SMR): the observation that in some colonies a large share of mites leave no adult daughter. The USDA laboratory at Baton Rouge selected first on this measure, then found that its main mechanism was the removal of infested pupae, and renamed the trait VSH.

What the measurements show

According to eXtension, in VSH colonies only about 30% of mites reproduce successfully and about 55% are infertile or lay no eggs. VSH bees respond strongly to heavily infested brood (15 to 25 mites per 100 capped cells) and much less to light infestations (1 to 5 per 100). They mostly remove unpigmented or lightly pigmented pupae (stages 2 to 4) and are much less hygienic towards infested drone brood. Recapping of opened cells reached 38% in VSH colonies, against 19% in hybrids and 17% in controls, in work by Villa and colleagues in 2010. Hybrid VSH colonies, with VSH queens open mated to unselected drones, keep significant resistance but less than the pure line.

Survivors: the Gotland experiment

Some European populations have survived untreated. On the Swedish island of Gotland, colonies left without treatment developed reduced mite reproduction. Barbara Locke, of the Swedish University of Agricultural Sciences, inseminated queens with single drones to make reciprocal crosses between Gotland and susceptible Uppsala bees. Reported in Apidologie in 2016, the share of mites reproducing was 0.43 in Gotland queen × Uppsala drone colonies against 0.85 in Uppsala × Uppsala colonies. The author concluded that the trait has a dominant genetic component rather than a maternal effect, but the study rested on four colonies per cross, and poor acceptance of single-drone queens limited replication.

The honest overall picture

The 2020 review by Matthieu Guichard and colleagues at Agroscope, in Genetics Selection Evolution, after three decades of selection programmes, concludes that the varroa problem has not been solved globally and that selection has not yet improved the survival of untreated colonies on a broad scale. The expression of these traits depends heavily on the environment: in one study, mite removal by VSH doubled after colonies were fed sugar syrup, and temperature, humidity, season, forage and brood quantity all change the measurements.

The UK government takes the same view. Answering a written question from Anna Gelderd MP on 25 July 2025, the Defra minister Mary Creagh said that developing varroa-tolerant bees through selective breeding is likely to be very difficult, because preserving resistance traits would need geographic isolation or artificial insemination and large numbers of colonies to keep genetic diversity. No government-funded breeding programme was named; beekeepers were pointed to BeeBase for varroa and queen rearing advice.

Hygienic queens in Britain

At the University of Sussex, Francis Ratnieks’s Laboratory of Apiculture and Social Insects (LASI) tested large numbers of hives for fast removal of dead brood, and in 2016 its spin-off began selling open mated and virgin hygienic queens to UK beekeepers. The university reported that hygienic hives had less than half the annual varroa build-up of other colonies and deformed wing virus levels 10,000 times lower. Open mated daughters will carry only half their genes from the selected mother, so the trait needs to be maintained by continued selection.

Whatever queens you buy, keep counting mites and use a medicine authorised by the Veterinary Medicines Directorate when the count calls for it. Our guide to varroa mites gives the thresholds.

What does genetics say about the British black bee?

It survived the imports

For decades British beekeepers were told that “Isle of Wight disease” killed off the native bee early in the twentieth century. Whole genome data now say otherwise. Buswell and colleagues, in the Journal of Apicultural Research in 2024, sequenced 70 colonies from Britain and Ireland. Seventeen showed no significant introgression from other subspecies. Colonsay and parts of South West England held colonies the authors describe as pure, and Ireland holds a large reservoir of A. m. mellifera.

Why crosses turn defensive

A first cross between distinct subspecies, such as a Carniolan or Buckfast queen mated to local dark drones, often shows hybrid vigour. When that cross’s daughters mate locally, the combinations that made it work are broken up and reshuffled. Some colonies inherit traits that work badly together, and temper is the one beekeepers notice first. Defensive behaviour has one of the higher heritabilities in the Swiss data, so a bad tempered drone source spreads quickly through an area. The cure is the same whatever the ancestry: requeen from a calm line and stop raising drones from the colony. Our page on honey bee breeds compares the bees kept in Britain.

Imports and the gene pool

Every imported queen adds new genes to the local drone population. Imports into Great Britain are legal but restricted. According to the NBU, only queens, each with no more than 20 attendants, may come from the EU; packages, nucs and colonies are banned. Every import must be notified on IPAFFS with an official health certificate, and within five days the original cage, attendants and packaging must be sent for examination, to the import address the NBU gives at the York Biotech Campus, Sand Hutton, for England and Wales, or to SASA in Edinburgh for Scotland. Northern Ireland is covered by DAERA, and only queens may be sent there from Great Britain, as an export with an official veterinary inspection. Whatever you decide about imported stock, register on BeeBase so inspectors can warn you of disease nearby.

Why do so many selection efforts fail?

Cause Mechanism Remedy
Open mating local drones supply half the workers’ genes and dilute the trait isolated sites, insemination, drone flooding
Low or misjudged heritability colony performance depends mostly on environment test many sister colonies, compare within the same apiary and year
Noisy mite counts drifting, robbing, brood levels, sampling traits measured in the cell (VSH, SMR), repeated counts
Environmental effects syrup, season, humidity and pesticides change VSH expression standard protocols, testing in several apiaries
Breeding populations too small loss of csd alleles, patchy brood, inbreeding many unrelated breeder queens, sharing stock within a group
Test versus real life a trait strong in tests may not keep a colony alive survival records over several winters
Mite adaptation a single resistance mechanism could select mites that bypass it combine traits, keep monitoring
Introgression imported queens mix local populations local stock, improvement groups, reserves such as Colonsay

How can a hobby beekeeper improve the genetics of their apiary?

You do not need an insemination kit to make progress. Here is a realistic approach for an apiary of a few National or 14x12 hives:

  1. Score every colony every year, the same way. Temper, calmness on the comb, swarming, spring build up, honey yield in kg or lb, winter stores used, and a mite count on a fixed date by natural drop or alcohol wash. Compare colonies in the same apiary, in the same year, with queens of the same age.
  2. Test hygienic behaviour on your best colonies with the NBU freeze-killed brood method. A domestic freezer is all you need. Always use the same reading time.
  3. Rear from two or three colonies only, the best over at least two seasons, and requeen the worst. Our guide to queen rearing gives the timetable.
  4. Flood your area with good drones. Leave drone comb in your breeder and drone colonies from spring and remove it from poor ones. Start about six weeks before you expect virgin queens to fly.
  5. Join a bee improvement group. BIBBA reckons a group needs at least 10 to 20 full colonies to start. Pooling records and drones gives a population that no single hobbyist can reach.
  6. Do not breed from a sick colony. Never graft from a colony showing any sign of notifiable disease, and report suspicion to your bee inspector at once.
  7. Widen your base if brood turns patchy without any disease: it often points to too narrow a population and matched csd alleles. A queen from an unrelated local breeder adds diversity.

What it costs

As listed in October 2026, an open mated, UK bred queen costs around £50, for example £50 including VAT for a British bred Buckfast queen from Bishops Bees, with delivery on top. Instrumentally inseminated breeder queens cost considerably more. For a small apiary, rearing a dozen queens a year from your own best colonies costs little more than a grafting tool, some cups and a few mating nucs, and keeps the genetics adapted to your local oil seed rape, June gap and, in the north and Scotland, the August heather.

Genetics will not replace good husbandry

A queen from a hygienic or VSH line lowers the risk of mite problems; it does not remove it. The colony still shares drones, drifting bees and robbers with every apiary around it. Treat breeding as a long-term investment that works best alongside regular mite monitoring, good nutrition and swift reporting of disease. The articles in our breeding section cover each of these techniques in more detail.

Frequently asked questions

Why does a drone have no father?

A drone hatches from an unfertilised egg, so it carries only one set of 16 chromosomes, all from its mother. Sex in honey bees is set by the csd gene: one copy gives a male, two different copies give a female. A drone therefore has a mother and a maternal grandfather but no father, and every sperm it produces carries an identical copy of its whole genome.

Can a patchy brood pattern be genetic?

Yes. If a queen mates with a drone carrying one of her two csd alleles, half the eggs fertilised by that drone's sperm are diploid males, which workers remove as young larvae. The empty cells give a scattered pattern. Inbreeding makes it more likely. Rule out an old or failing queen and brood disease first, and call your bee inspector if you see any sign of foulbrood.

Is hygienic behaviour inherited?

Partly, and it responds to selection better than most colony traits. In two Swiss populations its heritability was estimated at 0.06 to 0.19, and in the large European Carnica database it is the trait whose breeding values best predict later performance. The NBU's freeze-killed brood test is the standard way to score it. Mating must be controlled, or local drones dilute it within a generation.

Are there varroa resistant bees for sale in the UK?

Some UK breeders sell queens from colonies selected for hygienic behaviour, and a University of Sussex spin-off has sold hygienic queens since 2016. None is guaranteed to survive untreated. Defra told Parliament in July 2025 that breeding varroa-tolerant bees is likely to be very difficult. Keep monitoring mite levels and treat with a VMD authorised medicine when counts call for it.

What is the heritability of honey yield in bees?

Low to moderate, and it varies with the population and method. A Swiss study in Apidologie in 2020 estimated 0.02 to 0.11, while the European Carnica programme run through the BeeBreed database has made steady gains of about 336 g of honey a year since 2009. Low heritability means you need many related colonies, scored the same way, to select reliably.

Do I need instrumental insemination to improve my bees?

No. Most hobby beekeepers improve their stock by selecting carefully on records, rearing from their best two or three colonies and flooding the area with drones from good colonies. Insemination gives complete control of the father and is used by breeders and researchers, but it needs training, special equipment and enough colonies to avoid inbreeding. Joining a local bee improvement group is often more effective.

How far do drones fly to mate with a queen?

Further than most beekeepers expect. DNA paternity work in the Peak District found half of matings within 2.5 km and 90% within 7.5 km of the mating apiary, with one queen mating about 15 km away. A ridge between two valleys gave only partial isolation. That is why drones from every apiary within several kilometres shape the bees you raise.

Can I keep any honey bee I like in the UK?

Almost everywhere, yes. The exception is Colonsay and Oronsay in Scotland, where the Bee Keeping (Colonsay and Oronsay) Order 2013 has made it an offence since 1 January 2014 to keep any honey bee other than Apis mellifera mellifera. Elsewhere, imported queens must meet the import rules: from the EU, only queens with up to 20 attendants, notified on IPAFFS, with the cage sent for examination.

Sources

  1. European honey bee, Apis mellifera Linnaeus and subspecies (EENY-568/IN1005), University of Florida, IFAS Extension
  2. Sex determiner gene of honey bee more complicated than previously assumed (Lechner et al., Molecular Biology and Evolution, 2013), Phys.org, from the University of Cologne
  3. Discovery of the csd sex determination gene in the honey bee (Beye et al., Cell, 2003), news release, ScienceDaily, from the University of California, Davis
  4. An answer to the mystery of wanton queen honeybees: promiscuity produces more productive colonies (Mattila and Seeley, Science, 2007), Cornell Chronicle, Cornell University
  5. Estimates of genetic parameters for production, behaviour, and health traits in two Swiss honey bee populations (Apidologie, 51, 876-891, 2020), M. Guichard et al., Springer
  6. Substantial genetic progress in the international Apis mellifera carnica population since the implementation of genetic evaluation (Insects, 11, 768, 2020), A. Hoppe et al., MDPI
  7. Heritability in honey bees of characteristics associated with resistance to Varroa jacobsoni (Journal of Economic Entomology, 92(2), 261-265, 1999), J. R. Harbo and J. W. Harris, Oxford University Press
  8. Varroa sensitive hygiene and mite reproduction, Bee Health, eXtension (US land-grant university extension network)
  9. Advances and perspectives in selecting resistance traits against the parasitic mite Varroa destructor in honey bees (Genetics Selection Evolution, 52, 2020), M. Guichard, V. Dietemann, M. Neuditschko and B. Dainat, Springer
  10. Inheritance of reduced Varroa mite reproductive success in reciprocal crosses of mite-resistant and mite-susceptible honey bees (Apidologie, 47, 583-588, 2016), B. Locke, Springer
  11. Hygienic behaviour (Bee Craft, April 2002), National Bee Unit, APHA
  12. University of Sussex starts selling disease-resistant hygienic honey bee queens to UK beekeepers (2016), University of Sussex
  13. Bees: animal breeding, written question 69138, answered 25 July 2025, UK Parliament, written questions and answers
  14. Whole genome analyses of introgression in British and Irish Apis mellifera mellifera (Journal of Apicultural Research, 2024), Buswell et al., James Hutton Institute
  15. Quantifying honey bee mating range and isolation in semi-isolated valleys by DNA microsatellite paternity analysis (Conservation Genetics, 2005), Jensen et al., Royal Holloway repository
  16. The Bee Keeping (Colonsay and Oronsay) Order 2013 (SSI 2013/279), legislation.gov.uk
  17. BIBBA guide to forming and running bee improvement groups, Bee Improvement and Bee Breeders Association (BIBBA)
  18. Imports and exports, National Bee Unit, APHA (BeeBase)
  19. Honey bee health guidance: inspections, Scottish Government
  20. British bred Buckfast queen, open mated (price checked October 2026), Bishops Bees