KEEPA – BEE NETWORKING

Knowledge · The Bee Network

Know the animal
you keep.

Most beekeeping advice was written for European bees in European weather, and African bees do not read it. This is the underlying biology instead: how a bee is built, what she can sense that you cannot, how she finds her way home, and how a colony reaches a decision with nobody in charge. Every claim on this page carries the source it came from, and where the science disagrees with itself we say so rather than picking the tidier number.

01

Six numbers to start with

Bees are stranger than the folklore about them, and the strangeness is measurable.

20,000+Species of bee described worldwide. Seven of them make honey a person can harvest.
5,432Facets in one compound eye of a worker, each a separate light guide.
230HzWing beats a second, driven by a thorax that resonates rather than by nerve impulses.
960,000Neurons running the whole animal, in about a cubic millimetre of brain.
13.7kmThe longest foraging distance ever decoded from a waggle dance.
34.5°CThe temperature the brood nest is held at, whatever the weather outside.

03

How a bee is built

A honey bee is three boxes and a set of tools. The head carries the senses and the mouthparts, the thorax is almost entirely engine, and the abdomen holds the crop, the wax glands and the sting. Nothing on her is general purpose. The notch on her foreleg exists to clean an antenna, the hollow on her hind leg exists to carry pollen, and the hooks on her hind wing exist so that two wings behave as one.

Touch a label to read what the part does.

A worker honey bee in left side view with every external part named 12 to 15 mm, worker body length Three ocelli Forewing and hind wing Hamuli Compound eye Antenna Proboscis Thorax Corbicula Propodeum Spiracle Metasoma Sting
The worker, in fullTwelve to fifteen millimetres of animal. Choose a label to read what a part is for.
The worker honey bee in left side view, drawn to measured proportions. A queen and a drone differ in the ways set out in the table below.Chapter 2 of the Keepa bee reference. BEE-IMG-001.

The three castes are the same animal built to three specifications. A queen is longer and has no pollen basket because she will never forage. A drone is mostly eye and flight muscle, because his entire job is to find a queen in open air. A worker carries the full toolkit.

QueenWorkerDrone
Body length18 to 20 mm12 to 15 mm15 to 17 mm
Egg to emergence16 days21 days24 days
Lifespan2 to 7 years2 to 6 weeks in summerabout 90 days
Eye facets3,000 to 4,000about 5,4327,000 to 8,600
StingSmooth, reusable, kept for rival queensBarbed, torn out in mammal skinNone
Pollen basketNoneOn both hind legsNone
Ovaries120 to 200 ovarioles per ovary2 to 12, suppressedNone
What she doesLays, and holds the colony together chemicallyEverything elseMates once, then dies

04

What she can sense that you cannot

A bee works from a set of channels that overlap almost nowhere with ours. She sees ultraviolet and is blind to red. She reads the polarisation pattern of the sky, which to us is a blank blue. She detects the electric field a flower carries, feels the 265 Hz pulse of a dance through the comb in total darkness, and keeps a clock accurate enough to learn that a particular flower gives nectar at ten in the morning.

The eight channels a honey bee navigates and senses with, arranged around a bee at the centre Sun compassazimuth, corrected 15° an hour Polarised lightdorsal rim area, one blue patch is enough Optic flowdistance measured in image motion Smell170 odorant receptor genes Electric fieldshair deflection, Johnston's organ Magnetic fieldiron granules, threshold near 26 nT Vibration265 Hz dance pulses through comb Timea circadian clock that learns feeding hours
Eight channels a honey bee navigates and senses with. The ring lengths are drawn for legibility and carry no scale.Chapter 3 of the Keepa bee reference. BEE-IMG-015 and BEE-IMG-011.

Colour is the clearest case. Her three photoreceptors peak near 344, 436 and 544 nanometres, which shifts her whole window down the spectrum. She gains an ultraviolet channel and loses red. Many flowers exploit this with an ultraviolet absorbing centre against a reflecting rim, a landing target we cannot see at all.

The same flower as a person sees it and as a bee sees it, with the visible spectrum of each below AS A PERSON SEES IT Wavelength, nanometres 300 370 440 510 580 650 344 nm 436 nm 544 nm A PERSON SEES 380 TO 700 NM A BEE SEES ROUGHLY 300 TO 650 NM red, effectively invisible
The same flower to each of us, and the two visible ranges compared. The ultraviolet pattern is drawn in amber because ultraviolet has no colour a screen can show.Chapter 3 of the Keepa bee reference. BEE-IMG-009 and BEE-IMG-010.

06

Flight, and the myth about it

The claim that aerodynamics proves a bee cannot fly comes from Antoine Magnan in 1934, who applied fixed wing aircraft theory to a flapping wing and said in the same book that the failure must be in the theory. It was. Lift comes from leading edge vortices, from rotational lift as the wing flips at the end of each stroke, and from the wing recapturing energy in its own wake.

MeasureValueNote
Wing beatabout 230 HzFalls to about 211 Hz at 40 degrees Celsius, which sheds heat
Stroke amplitudeabout 90 degreesShort and fast, not long and slow
Cruising speed6 to 7 m/s, roughly 24 km/hSlower loaded, and the power curve bottoms near 3 m/s
Typical foraging range1 to 3 kmWhere a colony makes its living
Median danced distance6.1 km in an August dearthFrom 444 decoded dances
Longest decoded dance13.7 kmThe far edge of what a colony will do
Load carriedup to about 42 mgMeasured in the laboratory, near 60 per cent of body mass
Body temperature to flyabout 35 degrees CelsiusShe shivers to reach it before takeoff

Range is the number that matters to a keeper. A colony makes its living inside a kilometre or two and will go much further when it has to, which is why what grows near your stand matters more than what grows on the property line.

How far a colony forages, drawn as rings around the hive 1 kmMost trips. Where a colony makes its living. 3 kmThe working radius most beekeepers plan around. 6.1 kmMedian distance danced in an August dearth. 13.7 kmThe longest distance ever decoded from a dance. A COLONY MAY COVER 100 SQUARE KILOMETRES IN A SEASON
Foraging distance around one hive. Most of the work happens inside the first ring.Chapter 4 of the Keepa bee reference. BEE-IMG-021.

07

The colony decides, not the queen

The queen gives no orders. She lays, and she releases a pheromone that tells the colony she is present and laying. Everything else is decided from the bottom: which flowers get worked, when to build comb, when to raise a new queen, and where a swarm will live. A swarm choosing a cavity sends out scouts, argues by dancing, and reaches a quorum. It usually picks the best site available.

Caste is not genetic. A fertilised egg becomes a queen or a worker depending entirely on how it is fed in the first days of larval life. The same egg, fed differently, becomes a different animal with a different body and a lifespan fifty times longer.

Egg to adult for the queen, the worker and the drone on one time axis in days DAY 0 6 12 18 24 Queen emerges day 16 EGG 3 d LARVA 5.5 d CAPPED 7.5 d Worker emerges day 21 EGG 3 d LARVA 6 d CAPPED 12 d Drone emerges day 24 EGG 3 d LARVA 6.5 d CAPPED 14.5 d CELL CAPPED
The same fertilised egg becomes a queen or a worker. The difference is what it is fed, and it shows up as five days on the calendar and years on the lifespan.Chapter 5 of the Keepa bee reference. BEE-IMG-023.

08

Nectar to sealed cell

Honey is nectar that has been chemically altered and then dried. Invertase from the bee splits sucrose into glucose and fructose. Glucose oxidase produces gluconic acid, which takes the pH down near 3.9, and hydrogen peroxide, which keeps being produced whenever honey is diluted. Then the water goes, from somewhere near three quarters down to under a fifth, at which point nothing microbial can grow in it.

Nectar becoming capped honey in six stages, with what changes at each Nectar01 Sucrose, 70 to 80 per cent water, taken from the flower The crop02 About 40 mg carried home, invertase already working Handover03 Passed mouth to mouth to a receiver bee Tongue lashing04 Spread in thin films, water driven off Fanning05 Moved into a cell, ventilated below 20 per cent water Capped06 Sealed under wax, stable indefinitely 78% 78% 70% 45% 25% 18% WATER CONTENT
Six stages from the flower to a sealed cell. The bars beneath are the water leaving.Chapter 6 of the Keepa bee reference. BEE-IMG-032.
ComponentShareWhat it does
Fructoseabout 38 per centThe sweeter of the two sugars, and the reason honey stays liquid
Glucoseabout 31 per centCrystallises first, so a high glucose honey sets hard
Waterunder 20 per centBelow this figure nothing microbial can grow
Other sugarsabout 9 per centMaltose and higher sugars formed in the bee
Acids, minerals, enzymesabout 2 per centGluconic acid takes honey to a pH near 3.9
Hydrogen peroxideTrace, made continuouslyGlucose oxidase keeps producing it as honey is diluted

The distance behind a jar is the fact everyone repeats and almost everyone gets wrong. The commonly quoted figure is more than fifty thousand miles per pound, which is about 80,467 kilometres, or roughly 88,700 kilometres for a 500 g jar. Working it up from the other end, from crop capacity and round trip distance, gives a considerably larger number. The two do not agree, and we have left the disagreement visible rather than quietly picking one.

09

Two hundred facts

Every card carries a link to where the claim came from. Nothing here is folklore, and the things that turned out to be folklore are in the next section.

200 facts, every one with a source

The sensesFACT-001

A bee sees ultraviolet and is blind to red

Honey bee photoreceptors peak near 344, 436 and 544 nm, giving a visible range of roughly 300 to 650 nm. She gains a whole ultraviolet channel and loses red, so a pure red flower reads to her as a dark object.

J Comp Physiol A, 2014

The sensesFACT-002

Bees tell apart colours 4.5 nm apart

A honey bee discriminates wavelength differences as small as 4.5 nm with 70 per cent accuracy, near 400 nm and again near 500 nm. That is finer colour discrimination than most people assume any insect has.

J Comp Physiol A, 2014

The sensesFACT-003

Flowers wear targets we cannot see

Ultraviolet absorbing pigment at the centre of a petal against an ultraviolet reflecting rim makes a bullseye that is invisible to humans and unmissable to a bee. It works as a landing target.

NC State University, 2024

The sensesFACT-004

One worker eye holds 5,432 lenses

A worker compound eye averages 5,432 ommatidia, each sampling its own narrow cone of the world. A drone carries 7,000 to 8,600 per eye, and his meet across the top of his head.

Seidl and Kaiser, J Comp Physiol A

The sensesFACT-005

Bee eyesight is 200 times coarser than ours

Bee ommatidia sample the world about 1.7 degrees apart. A human fovea samples at about 0.008 degrees. The bee traded spatial resolution away and bought speed with the proceeds.

Seidl and Kaiser, J Comp Physiol A

The sensesFACT-006

A bee can see a flicker at 200 Hz

Honey bee flicker fusion has been measured at 200 Hz in bright light and 144 Hz in dim, against roughly 60 Hz for a human. A cinema screen would look like a slideshow to her.

PLOS ONE, 2014

The sensesFACT-007

A drone carries 30,000 smell plates per antenna

Olfactory pore plates number about 1,600 to 3,000 per antenna in a queen and 2,400 to 6,000 in a worker, but more than 30,000 in a drone. His entire investment is finding a queen at range.

apisavoir

The sensesFACT-008

170 genes for smell, 10 for taste

The honey bee genome carries 170 odorant receptor genes against 62 in the fruit fly, but only 10 intact gustatory receptor genes against 68. Smell is her precision sense and taste is nearly vestigial.

Genome Research

The sensesFACT-009

A bee tastes with her feet, and badly

Sugar triggers the feeding reflex at about 2.85 per cent on the antennae but needs about 34.23 per cent on the tarsi. No bitter receptor cells were found at the antennal tip at all.

Bee Culture, on de Brito Sanchez

The sensesFACT-010

Bees have no ears and still read the dance

Bees have no tympanal ear and cannot hear pressure waves. They detect the movement of air particles close to a vibrating source, using Johnston's organ inside the second antennal segment.

PLOS ONE, 2007

The sensesFACT-011

The dance can only be read within 3.5 mm

The effective detection zone around a waggle dancer is about 3.5 mm, so a follower must crowd in with her antennae almost touching. The dance is a private, physical signal, not a broadcast.

PLOS ONE, 2007

The sensesFACT-012

A 20 nanometre nudge is enough to register

The smallest antennal tip movement a bee detects is about 20 nanometres, driven by roughly 60 nanometres of air particle displacement. That is a few hundred atoms of travel.

PLOS ONE, 2007

The sensesFACT-013

A flying bee carries up to 450 volts

Friction with air and dust charges a bee's waxy cuticle positively, with reported potentials up to about 450 volts. Pollen jumps the gap to her from an earthed flower without contact.

National Geographic, 2013

The sensesFACT-014

Flowers carry electric fields and bees read them

Bumblebees detect and discriminate the electric fields of flowers, and learn two colours apart faster when electrical information is also available. The field is a real channel, not a side effect.

Science, 2013

The sensesFACT-015

A visited flower stays electrically marked

When a bee lands, the flower's electrical potential changes and stays changed for several minutes. In principle an approaching forager can tell a flower has just been emptied before wasting the landing.

University of Bristol, 2013

The sensesFACT-016

Bees carry magnetite in the abdomen

Iron rich granules about 0.3 micrometres across, holding magnetite nanoparticles of roughly 9.7 nanometres, sit in the fat body cells lining a bee's abdomen. How the signal reaches a neuron is still unsolved.

PLOS ONE, 2007

The sensesFACT-017

Cut one nerve and the magnetic sense stops

Bees trained to a 200 microtesla magnetic stimulus reached 70 per cent success by the 37th trial. Cutting the nerve cord between abdomen and thorax abolished that response while leaving smell intact.

Scientific Reports, 2016

The sensesFACT-018

A strip of eye reads the polarised sky

Along the top rim of each eye sits a band of specialised ommatidia whose optics deliberately blur the landscape so that only the polarisation pattern of the sky survives. It is a dedicated compass.

Frontiers in Cellular Neuroscience, 2018

The sensesFACT-019

One patch of blue sky is a whole compass

Rotate a polarising sheet over a dancing bee and her dance angle rotates with it. She needs one clear patch of sky to find the sun, and that patch need not contain the sun.

Nature, 1986

The sensesFACT-020

Bees keep five or six appointments a day

Trained to a feeder open only between set hours, bees return at that hour on later days with no food present. They hold about five or six daily time memories routinely, and up to nine with training.

J Comp Physiol A

Navigation and flightFACT-021

Bees measure distance in image motion, not metres

A bee's odometer counts the total image motion that swept across her eyes on the way out. It does not count metres, energy burned or wingbeats, which is why it can be fooled outright.

Nature, 2001

Navigation and flightFACT-022

A 6 m tunnel makes a bee report 184 m

Flown to a feeder 41 m away through a narrow textured tunnel, bees came home and danced for about 230 m. A feeder 6 m inside the tunnel was reported as about 184 m.

J Exp Biol, 2003

Navigation and flightFACT-023

Stripe the walls and the odometer stops turning

When tunnel walls were striped along the line of flight so they produced almost no image motion, 90 per cent of the returning dances reported under 50 m for a 41 m flight.

J Exp Biol, 2003

Navigation and flightFACT-024

Over water a bee underreports by three times

Bees trained across 240 m of open water in Canberra reported the distance honestly and wrongly, at roughly a third to a quarter of the truth, because featureless water generates almost no image motion.

PLOS Biology, 2004

Navigation and flightFACT-025

Sheffield bees flew a median of 6.1 km

Decoding 444 waggle dances at a Sheffield colony working heather in August gave a median foraging distance of 6.1 km, with some dances pointing beyond 9.5 km. In May the mean was about 1 km.

Beekman and Ratnieks, 2000

Navigation and flightFACT-026

Waggle dialects differ by species and place

Three Asian honey bee species compared in South India used different waggle durations for the same distance, and the species foraging furthest used the shortest run. Seconds per kilometre is a dialect, not a constant.

University of Würzburg, on Proceedings B 2020

Navigation and flightFACT-027

Displaced recruits flew the vector onto empty ground

Recruits that had followed a dance were caught leaving the hive, carried 200 to 250 m away and released. They flew the direction and distance they had been given, over ground where no feeder existed.

Riley et al., Nature, 2005

Navigation and flightFACT-028

Radar tracked bees correct for wind drift

The same harmonic radar study that ended the dance language argument also showed recruits adjusting their course mid flight to compensate for being blown sideways.

Riley et al., Nature, 2005

Navigation and flightFACT-029

Bees learn where the sun goes, they are not born knowing

Colonies moved to unfamiliar landscapes showed that bees acquire the relationship between the sun's daily course and the terrain they can see, then apply it in country they have never flown.

J Exp Biol, 2008

Navigation and flightFACT-030

Whether bees hold a map is still argued about

One camp reads homing from arbitrary release points as proof of a map like memory. The other says a large enough library of remembered views with attached vectors would look identical. It is unresolved.

PNAS, 2014

Navigation and flightFACT-031

Radar makes a bee answer on a different frequency

A batteryless transponder glued to the thorax reradiates a 9.4 GHz radar pulse at 18.8 GHz. Rocks, grass and buildings do not reply on the harmonic, so the bee stands out against silence.

Scientific Reports, 2019

Navigation and flightFACT-032

Wings beat 230 times a second through 90 degrees

A hovering honey bee uses an unusually short stroke at an unusually high frequency for an insect her size, the opposite of the pattern in most insects.

PNAS, 2005

Navigation and flightFACT-033

In thin air a bee widens her stroke by half

Flown in a low density helium and oxygen mixture, bees held wingbeat frequency almost constant and increased stroke amplitude by roughly 50 per cent. Frequency is nearly fixed; amplitude is the throttle.

PNAS, 2005

Navigation and flightFACT-034

Hovering costs 500 mW per gram of bee

Mass specific metabolic power in hovering flight runs at 500 ± 70 mW per gram, and bee flight muscle enzymes operate closer to their maximum catalytic capacity than any other muscle yet studied.

PNAS, 1996

Navigation and flightFACT-035

There is a genuinely economical cruising speed

Flight power against speed is a U shaped curve with a minimum near 3 m/s at about 37.5 mW, against 43 to 48 mW to hover. Flying slowly costs more than flying at a sensible clip.

J Comp Physiol B

Navigation and flightFACT-036

Water fails before heat does

In dry air at 40 °C a flying bee has about 1.5 hours before desiccation kills her, and just over 30 minutes at 46 °C. Her critical thermal maximum is 49 to 50 °C.

PNAS, 2024

Navigation and flightFACT-037

Alpine bumblebees hover higher than Everest

In a hypobaric chamber, alpine bumblebees hovered at air densities equivalent to about 7,900 m on average, and two individuals stayed airborne above 9,000 m, higher than the summit of Everest at 8,848 m.

Biology Letters, 2014

Navigation and flightFACT-038

A drone copied bee homing into 3.4 kB

A quadcopter running bee inspired path integration and visual homing in neural networks of 3.4 kB and 42.3 kB returned within 0.5 m of home on every flight of 30 to 110 m.

Nature, 2026

Honey and waxFACT-039

Nectar arrives four fifths water

Nectar reaching the hive is typically 70 to 80 per cent water. Finished honey is 15.5 to 18 per cent. Removing that water is the largest physical task in honey production.

Wikipedia, Honey

Honey and waxFACT-040

A 500 g jar is 33,000 to 50,000 crop loads

Working from a 40 mg crop at real arrival concentration, half a kilogram of honey takes between 33,000 and 50,000 full loads. The widely printed 25,000 assumes far richer nectar than most flowers give.

Codex Alimentarius honey standard

Honey and waxFACT-041

A forager makes 0.4 g of honey in her whole life

About one twelfth of a teaspoon is the lifetime output of one foraging bee. It is an order of magnitude estimate, and it is why a jar is a collective object.

SpaceDaily

Honey and waxFACT-042

A jar of honey is two to 50 million flower visits

Published flower counts per jar disagree wildly because nectar volume per flower ranges over three orders of magnitude between a lucerne floret and a eucalyptus blossom. Any single number is a species specific case sold as universal.

Wikipedia, Honey

Honey and waxFACT-043

Honey makes its own disinfectant when diluted

Glucose oxidase sits inactive in ripe honey for want of free water and switches on when honey is diluted, producing hydrogen peroxide at exactly the moment a microbial threat becomes real.

IntechOpen

Honey and waxFACT-044

The wax capping is the bees' own certificate

Workers seal a cell only once the contents are at or below roughly 18 to 20 per cent water. Uncapped stores can run as high as 70 per cent water and will ferment.

Wikipedia, Honey

Honey and waxFACT-045

Twenty minutes of tongue work per load

A receiver bee repeatedly spreads a droplet into a thin film between her mouthparts and reswallows it, for around 20 minutes per load. Thin film evaporation dries nectar far faster than a droplet sitting in a cell.

Wikipedia, Honey

Honey and waxFACT-046

1,100 wax scales make one gram

A wax scale is about 3 mm across and 0.1 mm thick and weighs roughly 0.9 mg. It sets in air after being secreted as a liquid through a smooth patch on the underside of the abdomen.

Beelistener

Honey and waxFACT-047

Wax costs up to 10 kg of honey per kilogram

Published conversion figures cluster between 6.6 and 10 kg of honey consumed per kilogram of wax secreted. Drawn comb is the most expensive thing a colony owns.

Wikipedia, Beeswax

Honey and waxFACT-048

One kilogram of wax carries 30 kg of honey

Against its build cost, comb is a spectacular structural bargain: a kilogram of beeswax holds 24 to 30 kilograms of honey.

Wikipedia, Beeswax

Honey and waxFACT-049

The hexagon saves six per cent of the wall

Among shapes that tile a plane with equal areas, a hexagon encloses a given area with roughly six per cent less perimeter than a square. Less perimeter means less wax.

Wikipedia, Honeycomb

Honey and waxFACT-050

Cells tilt 13 degrees so nectar cannot run out

Comb cells slope upward from the midrib by 9 to 14 degrees, commonly about 13. Ripening nectar is still liquid, and the tilt is what keeps it in the cell until capping.

Journal of Insect Science

Honey and waxFACT-051

Seven millimetres of gap invented modern beekeeping

Bees fill a gap wider than about 9.5 mm with comb and seal one narrower than 6.4 mm with propolis, leaving the space between open as a passage. Langstroth applied that clearance around every surface of a hanging frame in 1851.

Wikipedia, Langstroth hive

Honey and waxFACT-052

The movable frame was patented on 5 October 1852

United States Patent 9300 turned beekeeping from harvesting into husbandry. Before it, comb was fixed to the walls of a skep and taking honey usually meant killing the colony.

Wikipedia, Langstroth hive

Honey and waxFACT-053

Bees mummify what they cannot carry out

A mouse or lizard that dies inside a hive is too heavy to remove, so the bees encase the carcass in propolis. It is sealed, odourless and proof against decay.

Wikipedia, Propolis

Honey and waxFACT-054

Propolis is half resin and a third wax

Typical northern temperate propolis runs about 50 per cent resins and vegetable balsams, 30 per cent wax, 10 per cent essential oils and 5 per cent pollen, with roughly 50 constituents identified.

Wikipedia, Propolis

Honey and waxFACT-055

Bee bread is silage, and the pH proves it

Sealed pollen ferments through lactic acid bacteria: 72 hours at 35 °C dropped pH from 4.88 to 4.09 and lifted lactic acid from 3.46 to 4.82 per cent. Acidification is the preservation.

PMC10967301

Honey and waxFACT-056

Melittin is over half of bee venom

Melittin makes up about 52 per cent of the peptide fraction of apitoxin and is the compound that lyses cell membranes and causes the pain. Phospholipase A2, at about 12 per cent, is the principal allergen.

Wikipedia, Apitoxin

Honey and waxFACT-057

Honey crystallises fastest at 13 to 17 °C

Below 5 °C honey effectively will not crystallise, nucleation is most active at 5 to 8 °C, the fastest growth is at 13 to 17 °C, and crystals melt back at 40 to 50 °C. Granulation is a phase change, not spoilage.

Wikipedia, Honey

Honey and waxFACT-058

Manuka works through methylglyoxal, not peroxide

Most honey's antimicrobial punch comes from hydrogen peroxide. Manuka's comes chiefly from methylglyoxal, which is why it is graded separately, with 514 mg/kg of methylglyoxal corresponding to a UMF rating of 15.

Wikipedia, Manuka honey

Honey and waxFACT-059

Six times more manuka is sold than New Zealand makes

New Zealand produces roughly 1,700 tonnes of manuka honey a year. About six times that volume is marketed internationally as manuka, including an estimated 1,800 tonnes mislabelled in the United Kingdom alone.

Wikipedia, Manuka honey

Honey and waxFACT-060

Honey dressings close burns days faster

Preliminary clinical evidence indicates honey helps partial thickness burns heal four to five days faster than other dressings, and moderate evidence supports it for infected post operative wounds.

Wikipedia, Honey

The colonyFACT-061

A worker changes career five times in six weeks

She cleans cells for two days, nurses for nine, builds wax and receives nectar for six, guards and takes orientation flights for four, then forages until she dies. Her job list is set by which glands are currently working.

UF IFAS, ENY 166

The colonyFACT-062

The brood nest is steadier than a mammal

A colony holds its brood between 33 and 36 °C, targeting about 34.5 to 35 °C. Below roughly 32 °C pupae develop deformities.

PLOS ONE, 2010

The colonyFACT-063

Four degrees of brood heat change what a bee becomes

Brood reared at 32, 34.5 and 36 °C produced more than 4,400 tracked adults that differed measurably: the warmer reared bees were likelier to dance, foraged earlier and undertook more often.

Becher et al., 2009

The colonyFACT-064

Heater bees climb inside empty cells

Rather than warm the comb from the surface, heater bees press their thoraces against sealed cappings or climb into empty cells within the brood slab and heat the surrounding cells from inside.

PLOS ONE, 2010

The colonyFACT-065

A heater bee's thorax reaches 44.5 °C

Under extreme cold stress, heater bee thoracic temperatures up to 44.5 °C have been recorded, produced by shivering the flight muscles without moving the wings.

PLOS ONE, 2010

The colonyFACT-066

Bees can cook a hornet alive

Japanese honey bees engulf a hornet in a ball of 180 to 300 workers and shiver, holding the interior above 46 °C for as long as 20 minutes. The hornet dies; the bees, whose limit is a couple of degrees higher, do not.

Ono, Okada and Sasaki, Experientia, 1987

The colonyFACT-067

Bearding is cooling, not a symptom

When heat load rises, bees evacuate the cavity and hang outside the entrance, taking their own metabolic heat out of the nest and opening internal space for airflow. It is a functional response.

PLOS ONE, 2010

The colonyFACT-068

One sting leaves a pumping beacon behind

A worker's sting tears free with its venom sac, muscles and its own ganglion, and keeps sawing and pumping for minutes. The remnant also marks the target with alarm pheromone, recruiting more bees to that exact spot.

Wikipedia, Stinger

The colonyFACT-069

Alarm pheromone smells of banana

The principal sting alarm compound is isopentyl acetate, which smells strongly of banana. It is a good reason not to eat one next to an apiary.

NCBI Bookshelf

The colonyFACT-070

A bee bite is a local anaesthetic

Biting an intruder too small to sting, a bee injects 2-heptanone from her mandibles. Under a nanolitre paralyses a wax moth larva or a Varroa mite for a mean of 7.32 minutes.

PLOS ONE, 2012

The colonyFACT-071

A swarm takes three quarters of the workers

A primary swarm leaves with the old mated queen and 60 to 75 per cent of the workforce, weighted toward younger bees, plus most of the drones, all with their crops full of honey.

The Apiarist

The colonyFACT-072

Twenty to fifty scouts choose the new home

From the hanging bivouac, a small number of experienced bees search out candidate cavities and come back to dance for them on the surface of the cluster. Nobody else votes.

Wikipedia, Swarming

The colonyFACT-073

A swarm decides on quorum, not consensus

The move is triggered when 20 to 30 scouts are present at one site simultaneously. Lindauer's famous 1952 balcony swarm flew without ever reaching agreement, which is what pointed to a quorum in the first place.

Tom Seeley, Consensus or Quorum?

The colonyFACT-074

Scouts break a tie by silencing each other

Scouts committed to one site aim their stop signals at scouts dancing for the rival site, in proportion to their own numbers. That cross inhibition is what prevents a fatal split decision.

Cornell Chronicle, on Science 2012

The colonyFACT-075

A cavity under 15 litres is refused

Scouts assess against a demanding specification: at least about 15 litres, ideally 40, an entrance of about 12.5 square centimetres, low on the cavity, several metres up, dry, and preferably used by bees before.

Wikipedia, Swarming

The colonyFACT-076

Absconding is a strong colony's decision

Migrating African colonies studied in the Okavango were not weak ones fleeing. They had large populations, high brood production and full stores. Weak colonies were physically incapable of leaving.

Insectes Sociaux

The colonyFACT-077

A drone has a grandfather but no father

Drones develop from unfertilised eggs and are haploid, carrying 16 chromosomes to a female's 32. A drone's ancestor count in each preceding generation follows the Fibonacci sequence: 1, 1, 2, 3, 5, 8.

Wikipedia, Drone

The colonyFACT-078

Fewer than one drone in a thousand ever mates

A drone exists for one act he almost certainly will not perform. Those that do are killed by it: the endophallus everts explosively, tears away inside the queen and the drone falls and dies.

Wikipedia, Drone

The colonyFACT-079

25,000 drones gather at one spot in the sky

Drone congregation areas sit 10 to 40 m up and 30 to 200 m across, holding up to 25,000 drones drawn from as many as 200 colonies. They are aerial lekking sites.

Wikipedia, Drone

The colonyFACT-080

The same sky site is used for twelve years running

Drone congregation areas persist year after year, in one case with little change over 12 years, even though every drone using them this season died before last season ended. Nobody knows how they are found.

Wikipedia, Drone

The colonyFACT-081

Sisters share three quarters of their genes

Because a drone is haploid, every sperm he makes is identical, so full sisters share their father's entire genome. Relatedness between full sisters is 0.75, against 0.5 between a mother and daughter.

Wikipedia, Haplodiploidy

The colonyFACT-082

The textbook explanation of bee society is contested

The 0.75 figure only holds if a queen mates once. Real queens mate with 10 to 20 drones, dropping average relatedness to little above 0.25, and a recent analysis is titled "No evidence that haplodiploidy favors the evolution of eusociality".

PNAS

The colonyFACT-083

Diploid drones are eaten within hours

A fertilised egg homozygous at the sex determining locus becomes a diploid male, and workers detect and eat the larva almost immediately. The pepperpot pattern this leaves is a direct readout of inbreeding.

Nature Education

The colonyFACT-084

A larva has no working anus until the cell is capped

The larval hindgut is not connected to the stomach for the whole of larval life, because a larva lies directly in its own food. The connection opens only after capping, when feeding has stopped.

Bee Culture

The colonyFACT-085

A worker can hold her waste for four months

Six rectal pads reabsorb more than 90 per cent of the water in her waste, and the rectum is enormously expandable, letting a confined worker refrain from defecating right through a long winter.

Michigan State University

The colonyFACT-086

An adult bee never grows

A rigid exoskeleton cannot expand, so growth stops at the last moult. A small bee is not a young bee, she is a bee that was underfed as a larva. Adult size is a permanent record of larval nutrition.

Wikipedia, Insect morphology

The colonyFACT-087

Bees in the dark learn flowers they have never seen

Because nectar carries its floral scent through trophallaxis, bees deep inside the nest acquire olfactory memories of flowers they will only meet later, or never.

eLife, 2016

The colonyFACT-088

A forager who waits too long changes the subject

A returning forager who cannot find a receiver performs a tremble dance, which recruits more receivers and suppresses further departures. Unloading delay is the colony's internal price signal.

Wikipedia, Tremble dance

The colonyFACT-089

Foragers can send a location specific danger warning

Bees attacked at a feeder return and aim stop signals specifically at the bees advertising that source, not at dancers generally. It is a negative message about one place.

Frontiers, 2015

The colonyFACT-090

A chemical made in the honey crop delays adulthood

Ethyl oleate, produced in the crop of foraging bees and passed on by contact, delays young bees from starting to forage. The workforce regulates its own age profile chemically.

NCBI Bookshelf

The queenFACT-091

A queen is an ordinary egg fed differently

A queen and a worker start as the same fertilised egg. The difference is made entirely by diet and cell: a queen larva is fed royal jelly exclusively and without interruption for her whole larval life.

Wikipedia, Queen bee

The queenFACT-092

A queen emerges in 16 days, a worker in 21

Queens develop fastest of the three castes, at about 16 days from egg to emergence against 21 for a worker and 24 for a drone. Faster development is itself part of the queen advantage.

eXtension

The queenFACT-093

African queens emerge two days sooner still

Apis mellifera scutellata runs the whole schedule faster: 14 days to a queen against 16 in European bees, and 19 to 20 days to a worker against 21.

UF IFAS, EENY429

The queenFACT-094

A queen lays more than her own weight every day

At peak a good queen lays about 1,500 eggs a day, with exceptional queens recorded above 2,500. That is more than her own body mass daily, and around 200,000 eggs in a good year.

Wikipedia, Queen bee

The queenFACT-095

She mates in one week and never again

A virgin mates on warm afternoons six to ten days after emerging, at 10 to 40 m above the ground, and begins laying two to three days after her final return. The rest of her life runs on that one week.

Wikipedia, Queen bee

The queenFACT-096

Each mating lasts under five seconds

Copulation is often over in one or two seconds. The endophallus tears away and stays in the queen as the mating sign, which the next drone must displace.

Wikipedia, Drone

The queenFACT-097

Virgins toot and sealed queens quack back

A free virgin queen pipes with a two second pulse followed by quarter second toots. Queens still sealed in their cells answer with a muffled quacking. The exchange means an afterswarm may be imminent.

Drawwing, honey bee

The queenFACT-098

The first virgin out stings the others in their cells

A newly emerged virgin seeks out other emerged virgins and tears open sealed queen cells to sting the occupants. Only one survives, unless the colony is protecting cells for afterswarms.

Wikipedia, Queen bee

The queenFACT-099

The queen's sting is smooth and reusable

Unlike a worker's, a queen's sting is gently curved with much reduced barbs, so she can use it repeatedly and survive. She uses it almost exclusively on rival queens.

UF IFAS, EENY568

The queenFACT-100

Seventeen workers attend her at any moment

A retinue of about 17 to 18 workers surrounds the queen, licking, grooming and feeding her, then carries her pheromone out across the colony by contact, so every bee samples queen status continuously.

NCBI Bookshelf

The queenFACT-101

The queen leaves footprints that stop queen cells

Glands on the sixth segment of each of her legs deposit a footprint pheromone as she walks, which inhibits workers from building queen cups. Its decline is one of the first signs of a failing queen.

NCBI Bookshelf

The queenFACT-102

The queen reverses the usual rule of ageing

In most insects, high fertility costs longevity. The queen is simultaneously the most fecund and the longest lived member of the colony, which remains an unsolved problem in ageing biology.

Wikipedia, Queen bee

The queenFACT-103

The queen commands nothing

She does not allocate foragers, choose a nest site, decide when to swarm or direct defence. Every one of those is a distributed computation running on dances, stop signals and pheromones she does not produce.

NCBI Bookshelf

Numbers that do not seem realFACT-104

A bee brain runs on 960,000 neurons

The whole apparatus behind navigation, dance, learning and colony decision making holds roughly 960,000 neurons in about one cubic millimetre of tissue. A human brain has about 86 billion.

BioNumbers

Numbers that do not seem realFACT-105

170,000 memory cells per mushroom body

Each of the bee's paired mushroom bodies, the seat of olfactory learning and memory, holds about 170,000 densely packed Kenyon cells, against 2,500 in a fruit fly. That is roughly 68 times as many.

Learning and Memory

Numbers that do not seem realFACT-106

The wing pair locks together with hooks

A row of hooks along the leading edge of the hind wing catches into a fold on the trailing edge of the forewing, making one aerofoil in flight. At rest the hooks disengage and the wings fold separately.

Michigan State University

Numbers that do not seem realFACT-107

The world holds more than 20,000 bee species

Apis mellifera is one species out of over twenty thousand described, across seven families, and new ones are described every year. The living total is certainly higher than the tally.

Discover Life

Numbers that do not seem realFACT-108

Over ninety per cent of bees are solitary

Eusociality, the arrangement every beekeeper takes as normal, is the exception among bees. It has evolved independently several times, and most bee species have no workers, no queen and no hive.

Wikipedia, Eusociality

Numbers that do not seem realFACT-109

More than 3,000 bee species build nothing

Cuckoo bees gather no pollen, have no pollen carrying structures and construct no nest. The female enters a host nest and either she or her larva destroys the host egg and eats its provisions.

Britannica, cuckoo bee

Numbers that do not seem realFACT-110

One comb, one metre across, 45 kg of honey

Apis dorsata builds a single vertical comb up to a metre wide on cliffs and buildings, holding up to 100,000 bees and as much as 45 kg of honey. Up to 200 colonies have been recorded in one tree.

Wikipedia, *Apis dorsata*

Numbers that do not seem realFACT-111

Himalayan cliff bees nest at 3,000 m

Apis laboriosa, the largest honey bee at up to 30 mm, nests under overhangs at 2,500 to 3,000 m and forages to 4,100 m. Its spring honey carries a toxin and fetches about five times the ordinary price.

Wikipedia, *Apis laboriosa*

Numbers that do not seem realFACT-112

One hive eats what 100,000 wild bees would need

Over three months a single honey bee colony collects pollen equivalent to what would support around 100,000 native solitary bees. That is why hive placement in a nature reserve is a conservation cost.

Xerces Society

Numbers that do not seem realFACT-113

The famous €153 billion is value at risk, not loss

The most quoted pollination figure assumes the total disappearance of pollinators and ignores every adaptive response open to farmers. Its authors are explicit that it is not a forecast.

Helmholtz Centre UFZ

Numbers that do not seem realFACT-114

35 per cent of world crop volume depends on pollinators

The FAO puts three quarters of the world's most productive crop plants as at least partly pollinator dependent, over 80 per cent of flowering plant species as animal pollinated, and counts more than 25,000 bee species among about 200,000 pollinators.

FAO

Numbers that do not seem realFACT-115

Deformed wing virus rose a millionfold in Hawaii

When Varroa arrived in Hawaii, deformed wing virus prevalence went from about 10 per cent to 100 per cent, viral load rose around a millionfold, and strain diversity collapsed to one dominant variant.

Martin et al., Science, 2012

Numbers that do not seem realFACT-116

A mite feeds on the organ that runs immunity

For fifty years textbooks said Varroa drank bee blood. It does not: it feeds primarily on the fat body, the organ handling detoxification, immunity and overwintering reserves, which is why its damage compounds every other stressor.

Ramsey et al., PNAS, 2019

Numbers that do not seem realFACT-117

One small hive beetle female lays 2,000 eggs

Aethina tumida females lay 1,000 to 2,000 eggs, the sex ratio runs about 1.58 females per male, and wandering larvae travel over 200 m through soil to pupate. Adults live up to six months.

Journal of Integrated Pest Management, 2022

Bees and peopleFACT-118

The oldest picture of a honey hunt is 8,000 years old

At the Cuevas de la Araña near Valencia, a painted figure climbs a rope or grass ladder into a cliff cavity while bees circle. It is the earliest known depiction of bees and of people eating honey.

Wikipedia, Cuevas de la Araña

Bees and peopleFACT-119

Egypt kept bees before the pyramids at Giza

Egyptian beekeeping is recorded from roughly 3100 BCE, and a relief in the sun temple of Nyuserre Ini, carved before 2422 BCE, shows the whole sequence: smoking hives, removing combs, straining and sealing honey into jars.

Wikipedia, Beekeeping

Bees and peopleFACT-120

A 2,900 year old apiary made 500 kg a year

At Tel Rehov in Israel, dated to about 900 BCE, archaeologists found roughly thirty intact straw and clay cylinder hives in ordered rows, an industrial operation capable of around half a tonne of honey annually.

Wikipedia, Beekeeping

Bees and peopleFACT-121

Mead may be the oldest alcoholic drink of all

Pottery from northern China dating to at least 7000 BCE carries chemical signatures consistent with honey, rice and fermentation. Honey wine plausibly predates both beer and grape wine.

Wikipedia, Mead

Bees and peopleFACT-122

The bee was the hieroglyph for Lower Egypt

The pharaonic throne name used the title nesu bity, read as "he of the sedge and the bee". The sedge stood for Upper Egypt and the bee for Lower Egypt, so half the kingdom was written as an insect.

Wikipedia, Ancient Egyptian royal titulary

Bees and peopleFACT-123

A Frankish king was buried with 300 golden bees

The tomb of Childeric I, opened in 1653 at Tournai, held some 300 golden winged insects laid on the king's cloak. Most were stolen and melted down in 1831; two were later fished out of the Seine.

Wikipedia, Childeric I

Bees and peopleFACT-124

Napoleon took a dead king's bees as his emblem

In June 1804 the Conseil d'État approved using Childeric's bees as symbols of the French Empire, replacing the Capetian fleur de lys. Napoleon's coronation regalia is covered in them.

Wikipedia, Childeric I

Bees and peopleFACT-125

Deseret means honeybee, and Utah is the Beehive State

The word comes from the Book of Mormon and translates as honeybee. It named the provisional state, the University of Deseret and, through the hive emblem, the state's whole visual identity.

Wikipedia, Deseret

Bees and peopleFACT-126

The sixteenth chapter of the Quran is called The Bee

Surah An-Nahl has 128 verses and takes its name from verses 68 and 69, which describe bees building in hills, trees and human dwellings, and a drink of varying colours issuing from them in which there is healing.

Wikipedia, An-Nahl

Bees and peopleFACT-127

Xenophon's army was floored by honey in 401 BC

Greek soldiers near Trabzon ate local honey and, in Xenophon's account, those who ate a little were like drunk men and those who ate a lot behaved like madmen. All recovered the following day.

Wikipedia, Mad honey

Bees and peopleFACT-128

Honey was used as a weapon against Rome in 65 BCE

During the Third Mithridatic War, Mithridates VI had toxic honeycomb placed along Pompey's line of retreat. Strabo records that it wiped out three maniples of Roman soldiers, somewhere between 480 and 1,800 men.

Wikipedia, Mad honey

Bees and peopleFACT-129

Mad honey still poisons people, mostly in Turkey

Rhododendron nectar makes honey carrying grayanotoxin. In one analysis 91 per cent of poisoning cases came from the Black Sea region of Turkey and 5 per cent from Nepal, mostly in middle aged men seeking a perceived aphrodisiac.

Wikipedia, Mad honey

Bees and peopleFACT-130

Beekeepers still tell the bees about a death

The custom of informing hives of a death, birth or marriage is recorded across England, Ireland, Wales, Germany, the Netherlands, France, Switzerland, Bohemia and New England. Hives were draped in black and told in a low voice.

Wikipedia, Telling the bees

Bees and peopleFACT-131

The palace bees were told the Queen had died

In 2022 the royal beekeeper John Chapple went to the hives at Buckingham Palace and Clarence House to inform the bees of Elizabeth II's death and of the accession of Charles III.

Wikipedia, Telling the bees

Bees and peopleFACT-132

World Bee Day is a Slovenian beekeeper's baptism day

The United Nations adopted 20 May in December 2017, on a Slovenian proposal. The date is the day Anton Janša, the pioneer of modern beekeeping, was baptised in 1734.

Wikipedia, World Bee Day

Bees and peopleFACT-133

Sherlock Holmes retired to keep bees

Conan Doyle sent Holmes to a small farm on the Sussex Downs with beekeeping as his main occupation, and had him write a Practical Handbook of Bee Culture. It is the only career Holmes ever chose for himself.

Wikipedia, Sherlock Holmes

Bees and peopleFACT-134

The smoker was invented in 1873 and never patented

Moses Quinby built the modern bee smoker in the Mohawk Valley and, as a Quaker, deliberately declined to patent it, handing the design to beekeeping. Almost every smoker since is a variation on it.

Wikipedia, Bee smoker

Bees and peopleFACT-135

A wrecked galleon still leaves beeswax on an Oregon beach

The Spanish Manila galleon Santo Cristo de Burgos was lost in the winter of 1692 to 1693. Its cargo of beeswax has been washing ashore ever since, and the wings of Philippine bees are trapped inside it.

Wikipedia, Beeswax wreck

Bees and peopleFACT-136

A million hives are trucked to Californian almonds

Nearly half of all beehives in the United States are moved to the almond orchards each February. It is the largest annual managed pollination event in the world.

Wikipedia, Almond

Bees and peopleFACT-137

Honey is not safe for a baby under one year

Honey can carry Clostridium botulinum spores that an infant gut cannot handle. Infant botulism runs at about 1.9 cases per 100,000 live births in the United States, and honey is considered safe only after the first year.

Wikipedia, Honey

Bees and peopleFACT-138

Bee venom therapy has no evidence behind it

There is no scientific evidence that bee venom therapy treats or alters the course of cancer or any other disease. Trials in multiple sclerosis found it ineffective and capable of worsening symptoms.

Frontiers in Immunology, 2024

Bees and peopleFACT-139

The Schmidt sting pain index was built by being stung

Justin Schmidt published his scale of insect sting pain in 1983, rating stings from 0 to 4. The honey bee sits at level 2, described as an oven mitt with a hole in it when you pull the cookies out.

Wikipedia, Schmidt sting pain index

Bees and peopleFACT-140

Bees fell silent at the moment of totality

Acoustic monitoring across three regions during the 2017 total solar eclipse found that bees stopped flying entirely at totality, but flew normally through the dim partial phases. Light level, not temperature, was the trigger.

Annals of the Entomological Society of America

South AfricaFACT-141

South Africa has close to 1,000 bee species

The country holds roughly a thousand recorded bee species in about 80 genera, many endemic to the Fynbos and Succulent Karoo. Nearly a third of those genera still need taxonomic revision.

University of Cape Town News

South AfricaFACT-142

Two honey bee subspecies meet in the Karoo

South Africa is the only country where two honey bee subspecies with fundamentally different reproductive biology meet along a natural boundary, and where that meeting is a regulated agricultural matter.

SANBI

South AfricaFACT-143

The bee boundary is 200 km wide and written into law

The hybrid zone between the Cape bee and the savannah bee runs 100 to 200 km wide through the semi arid Karoo, between about 31°S and 33°S. Moving colonies across it is restricted in either direction.

Beaumont et al., Heredity

South AfricaFACT-144

Cape worker bees lay daughters without mating

Apis mellifera capensis workers reproduce by thelytokous parthenogenesis: an unfertilised worker laid egg develops into a diploid female. No other honey bee in the world does this.

Annals of the Entomological Society of America, 2021

South AfricaFACT-145

Four hundred hives moved north killed fifty thousand

Roughly 400 Cape colonies were carried into savannah bee territory in 1992. By 1993 an estimated 50,000 colonies had been destroyed. It is known as the capensis calamity.

Beaumont et al., Heredity

South AfricaFACT-146

One parasitic clone has run for over thirty years

The capensis social parasite persists as a clonal lineage: a single genetic individual reproducing asexually inside host colonies, passed from hive to hive, for more than three decades.

Annals of the Entomological Society of America, 2021

South AfricaFACT-147

A parasite bee mimics a queen and stops the foraging

A Cape worker in a savannah colony activates her ovaries, produces a queenlike pheromone blend and is fed as a queen. Her clonal daughters do the same, the foraging force collapses and the colony dies.

SANBI

South AfricaFACT-148

Fynbos holds over 9,000 plant species

The fynbos of the Cape carries more than 9,000 plant species, around 6,200 of them found nowhere else. It is the smallest and richest floral kingdom in the world per unit of area.

Wikipedia, Fynbos

South AfricaFACT-149

South Africa's flagship honey is slow to set

Fynbos honey is amber to dark, complex and slightly resinous, made from Cape Erica, Protea and Leucospermum, and its sugar profile keeps it liquid far longer than most South African honeys.

Agribook

South AfricaFACT-150

There is not one native bumblebee in South Africa

Bumblebees occur in Africa only north of the Sahara. Every buzz pollination service in South African agriculture is performed by carpenter bees, Amegilla, halictids and other native solitary bees.

Wikipedia, Bumblebee

South AfricaFACT-151

A South African bee has legs longer than its body

Rediviva females collect floral oil with forelegs running from 6.9 mm to 23.4 mm, in some species longer than the entire bee. All 26 species live only in southern Africa.

Kahnt et al.

South AfricaFACT-152

Here the flowers led and the bees followed

Sampling 19 of the 26 Rediviva species against the paired oil spurs of about 70 Diascia species showed the bee legs tracking the flowers rather than the reverse. It is one of the few documented cases of a pollinator adapting to the plant.

ScienceDaily

South AfricaFACT-153

Southern Africa has a stingless bee that nests underground

The mopane bee, Plebeina hildebrandti, is one of only three African stingless bees that nest below ground in grassland. It gives a dark, strongly flavoured honey and clusters at eyes and mouths for moisture in dry country.

icipe

South AfricaFACT-154

South Africa eats more than twice the honey it makes

Production is put at roughly 2,000 tonnes a year against about 5,000 tonnes consumed, with over 3,000 tonnes imported to close the gap. In 2024 the country imported R104 million of honey and exported R45 million.

Agribook

South AfricaFACT-155

The last national colony census was in 1975

Every circulating figure for how many managed colonies South Africa holds rests on a count taken half a century ago. The commonly quoted 200,000 is not attached to any survey.

Melin et al., SAJS 2014

South AfricaFACT-156

Wild insects pollinate more value than managed hives

In the South African deciduous fruit industry, managed honey bee pollination was valued at R46 million and wild insect pollination at R53 million. The unmanaged half of the workforce does more.

Melin et al., SAJS 2014

South AfricaFACT-157

The industry's main nectar source is a listed invader

South African natural vegetation is a poor bulk nectar source, and the industry leans heavily on introduced Eucalyptus. Six eucalyptus species are listed Category 1b invasive and must be controlled.

SANBI

South AfricaFACT-158

Macadamia orchards are still short of pollination

South African macadamia orchards have shown pollination limitation persisting despite managed honey bee stocking. Hive numbers alone do not guarantee that a crop is adequately pollinated.

Melin et al., SAJS 2014

South AfricaFACT-159

235 hives were destroyed in a single night

In late May 2021 a South African beekeeper lost 235 bee boxes in one night, valued at about R700,000. A separate theft took 60 hives from a sunflower farm at about R3,000 per super.

Daily Maverick, 2021

South AfricaFACT-160

Eighty per cent of surveyed beekeepers fight honey badgers

Research summarised for South Africa found over 80 per cent of surveyed beekeepers had honey badger problems and more than half admitted killing badgers. Raising hives a metre onto stands is the standard fix.

Science in Africa

South AfricaFACT-161

American foulbrood reached the Cape in December 2008

The first clinically symptomatic colonies appeared in the Western Cape in December 2008 and were confirmed in February 2009. An industry estimate put the loss at around 40 per cent of the Western Cape bee population.

The Conversation

South AfricaFACT-162

Varroa arrived in 1997 and is mostly left untreated

The mite reached the Western Cape in 1997 and spread across the country within a few years. South African beekeepers largely do not treat, and colony development in treated and untreated apiaries did not differ significantly in one season's comparison.

Strauss et al.

South AfricaFACT-163

Savannah bees suppress the mite themselves

African savannah honey bees show resistance rather than tolerance, actively suppressing mite population growth by grooming and by removing infested brood, helped by short post capping periods and frequent absconding.

Parasitology

South AfricaFACT-164

A bird leads people to bees, and it is only African

The greater honeyguide knows where wild colonies are but cannot open them. People can open a nest but cannot find one. In parts of Africa the two cooperate deliberately, and it is the only known case of its kind.

African Honeyguides

South AfricaFACT-165

The right call doubles the chance of being led to honey

Working with Yao honey hunters in Mozambique, a specialised trilled call raised the probability of being guided from about 33 to 66 per cent and of finding a nest from about 17 to 54 per cent against control sounds.

Spottiswoode, Begg and Begg, Science, 2016

South AfricaFACT-166

A honeyguide raises Hadza nest finding by 560 per cent

Among the Boran of Kenya, following a honeyguide cuts honey search time by about two thirds. In northern Tanzania, honeyguides raised the Hadza rate of finding nests by 560 per cent and led them to richer nests.

Wikipedia, Greater honeyguide

South AfricaFACT-167

The honeyguide chick kills its nest mates with a hook

A honeyguide nestling hatches blind and featherless with a membranous hook on its bill, which it uses to kill the host's young outright or by repeated wounds. The same species that helps people is a brood parasite.

Wikipedia, Greater honeyguide

South AfricaFACT-168

Beehive fences cut elephant raids by up to 85 per cent

Lucy King of Save the Elephants noticed that elephants avoid acacias holding bee colonies, and built fences of hives at chest height every 10 m. They reduce human elephant conflict by up to 85 per cent and produce honey.

Wikipedia, Beehive fence

South AfricaFACT-169

African bees send four times the defenders

Savannah bees and their Africanised descendants differ quantitatively, not qualitatively: they react faster, send three to four times as many workers to a disturbance, deliver about ten times as many stings and pursue up to 400 m.

Wikipedia, Africanized bee

South AfricaFACT-170

Twenty six swarms escaped in October 1957

A visiting beekeeper removed queen excluders from experimental hives near São Paulo holding African queens imported from southern Africa, and 26 swarms got out. Their descendants reached Texas by 1990.

Wikipedia, Africanized bee

Intelligence and learningFACT-171

Bees learn "same" and "different" as abstract rules

Trained in a Y maze to pick the stimulus matching a sample, or the one that does not, bees applied the rule to stimuli they had never seen. They had learned a relation, not a picture.

Nature, 2001

Intelligence and learningFACT-172

A rule learned by eye transfers to smell

Bees trained on the matching rule with colours applied it to patterns, and bees trained visually applied it to odours. The rule crossed sensory modalities intact.

Nature, 2001

Intelligence and learningFACT-173

Bees put zero at the low end of a number line

Trained on "choose fewer", bees shown an empty card chose it over one element 63 per cent of the time, and did better against five or six elements than against one. That numerical distance effect had previously been shown only in some primates and parrots.

Science, 2018

Intelligence and learningFACT-174

Bees recognise human faces

Bees trained on photographs of people could tell individuals apart and recognise them later. Nobody claims a face detection module; a face is probably processed as an oddly shaped flower.

J Exp Biol, 2005

Intelligence and learningFACT-175

Bumblebees improved on their teacher

Shown a demonstrator rolling the furthest ball to a target for sugar, observer bees rolled the nearest instead, and used balls of a colour never demonstrated. They took the point rather than the recipe.

QMUL, on Science 2017

Intelligence and learningFACT-176

No bee solved the puzzle box alone in 24 days

Across three colonies given 12 to 24 days, no bumblebee opened a two step puzzle box on its own, while 5 of 15 observers learned it from a demonstrator, three on the first attempt.

Nature, 2024

Intelligence and learningFACT-177

Bees sleep, and sleep loss ruins their dancing

Foragers deprived of sleep overnight danced with degraded directional precision the following day. Bees also consolidate navigation memory during sleep.

PNAS, 2010

Intelligence and learningFACT-178

Six pairings teach a bee to sting a smell

Aversive conditioning pairs an odour with a two second, 7.5 volt shock. Six pairings are enough for the odour alone to trigger the sting extension reflex.

PLOS ONE, 2007

Intelligence and learningFACT-179

The waggle dance is partly learned

Colonies of same age bees, where no young bee ever watched an experienced dancer, danced far more disorderly. The angular disorder healed over about 20 days.

Science, 2023

Intelligence and learningFACT-180

Identified neurons fire one spike per dance pulse

Interneurons in the bee's primary auditory centre track the dance pulse train, with inhibitory cells apparently encoding how long the train lasts, which is the variable carrying distance.

J Neuroscience, 2017

Intelligence and learningFACT-181

A swarm decides the way a brain decides

Scouts excite through dancing and inhibit through stop signals aimed at the rival camp, which is formally the cross inhibition that neural circuits use to turn competing evidence into one choice. Seeley called it analogous, and meant the word.

Cornell Chronicle

Intelligence and learningFACT-182

500 scientists signed for a realistic possibility

The 2024 New York Declaration on Animal Consciousness, signed by more than 500 scientists, states that the evidence indicates at least a realistic possibility of conscious experience in many invertebrates including insects.

Wikipedia, New York Declaration

The strange and the extremeFACT-183

The largest bee lives inside a termite nest

Megachile pluto, Wallace's giant bee, has a female body of 38 mm and a wingspan of 63.5 mm. She nests inside active termite mounds, partitioning the galleries with resin.

Wikipedia, *Megachile pluto*

The strange and the extremeFACT-184

The smallest bee weighs a third of a milligram

Perdita minima is just under 2 mm long and weighs about 0.33 mg. It nests in sandy soil in the deserts of the southwestern United States.

Wikipedia, *Perdita minima*

The strange and the extremeFACT-185

One bee's tongue is twice its own body length

The orchid bee Euglossa natesi, described from Colombia, has a tongue of 33.76 mm, roughly double its body length.

Mongabay

The strange and the extremeFACT-186

Three species of bee eat meat

The vulture bees, three Neotropical stingless species, are obligate necrophages. Foragers enter carcasses through the eye sockets, fill the crop with flesh and regurgitate it into storage pots. They keep no pollen at all.

Wikipedia, Vulture bee

The strange and the extremeFACT-187

A bee that builds its nursery out of flower petals

Osmia avosetta lines a shallow ground tunnel with flower petals glued together with mud, provisions the chamber and folds the petals over. It was documented simultaneously in Turkey and Iran in 2009.

Wikipedia, *Osmia avosetta*

The strange and the extremeFACT-188

Male orchid bees collect perfume, not pollen

Male euglossine bees spend their lives gathering volatile compounds from orchids, apparently as an advertisement of fitness. Whole orchid groups are pollinated exclusively by males, and the females never visit them.

Wikipedia, Euglossini

The strange and the extremeFACT-189

The bee louse is a wingless fly 1.6 mm long

Braula coeca is a fly so flattened and reduced that it is barely recognisable as one. Adults ride on the bodies of honey bees and the larvae tunnel through wax. Whether it does any harm is still argued.

Wikipedia, *Braula*

The strange and the extremeFACT-190

A fly turns honey bees into night flying zombies

Parasitised by Apocephalus borealis, bees abandon the hive at night, walk in circles, lose the ability to stand and gather at lights. The larvae feed on flight muscle and emerge about a week later.

Wikipedia, *Apocephalus borealis*

The strange and the extremeFACT-191

Asian honey bees paste dung round their door

Apis cerana colonies collect animal faeces and stick spots of it about 2 mm across around the nest entrance. Hornets spent 94 per cent less time trying to chew their way into a spotted entrance.

Wikipedia, *Vespa soror*

The strange and the extremeFACT-192

Giant honey bees defend with a stadium wave

Apis dorsata colonies run shimmering waves of hundreds of bees flipping their abdomens upward, complete in under 400 ms and involving up to 70 bees within 600 ms. Hornets react in under 100 ms.

PLOS ONE, 2008

The strange and the extremeFACT-193

An orchid that pretends to be a female bee

Ophrys apifera mimics both the scent and the appearance of a female bee, and male Eucera longicornis attempt to mate with the flower, picking up pollen in the process. In northern Europe the plant gave up and pollinates itself.

Wikipedia, *Ophrys apifera*

The strange and the extremeFACT-194

The oldest known bee is in New Jersey amber

Cretotrigona prisca, a stingless bee from the Late Cretaceous, is the earliest fossil bee known. The specimen appears to be a worker, so eusociality was already running before the dinosaurs ended.

Wikipedia, *Cretotrigona*

The strange and the extremeFACT-195

Bees are wasps that changed what they ate

Bees evolved from inside the apoid wasps. The switch was larval diet: prey dusted with pollen, then larvae raised partly on pollen, then no meat at all. Branched hairs, pollen baskets and the whole pollination economy follow from that.

Wikipedia, Bee

The strange and the extremeFACT-196

Bees are about 124 million years old

Phylogenetic dating places the origin of bees in the Early Cretaceous, roughly 124 million years ago, on the palaeocontinent of West Gondwana, alongside the early radiation of the flowering plants rather than after it.

Wikipedia, Bee

The strange and the extremeFACT-197

The reliable way to spot a fake bee is counting wings

Hoverflies mimic bees so well that colour and fuzz are useless. Flies have one pair of wings, very short antennae and enormous eyes meeting on top of the head, and they hover motionless then dart sideways.

Wisconsin Horticulture Extension

The strange and the extremeFACT-198

Sweat bees are after your salt

Halictid bees are named for their attraction to perspiration, which they visit for the dissolved electrolytes. Most stings from them happen when somebody swats one that is trying to drink.

Wikipedia, Halictidae

The strange and the extremeFACT-199

The alfalfa seed crop runs on a ground nesting bee

The alkali bee, Nomia melanderi, snaps open the lucerne flower keel in a way honey bees avoid. Growers build salty mud beds so it will nest, having learned that ploughing the natural flats cut seed yields dramatically.

Wikipedia, *Nomia melanderi*

The strange and the extremeFACT-200

A detached sting keeps working on its own

The torn out sting apparatus carries its own ganglion, so the lancets go on sawing and the valves go on pumping venom for several minutes after the bee that delivered it has gone.

Wikipedia, Stinger

10

What everybody knows, and what is true

These are the claims that came up again and again in the research and did not survive it. Several of them are repeated by beekeepers.

Aerodynamics proves a bee cannot fly

The claim traces to Antoine Magnan and a mathematician collaborator in 1934, who applied fixed wing theory to a flapping wing, and Magnan said in the same book that the failure was in the theory. Lift comes from leading edge vortices, rotational lift and wake capture. Wikipedia, Antoine Magnan

A bee dies whenever she stings anything

The barbs only lodge in elastic mammal skin. Against another insect, whose cuticle is not elastic, a worker stings repeatedly and survives, and a queen's sting is smooth and reusable. Wikipedia, Stinger

Honey never needs any care

Honey is stable only while it stays dry and sealed. Above roughly 20 per cent water osmophilic yeasts ferment it, and heat and age raise HMF, which is why the Codex standard sets limits for both. Codex Alimentarius honey standard

Edible honey has been recovered from Egyptian tombs

This is the most repeated honey claim in the world and it does not survive checking. No edible honey has been found in an Egyptian tomb; every such case has proved to be another substance or only chemical traces. Wikipedia, Honey

The queen gives the orders

Queen pheromone changes what workers do, but it is one signal inside a feedback system. Foraging effort is regulated by tremble dances, stop signals, brood pheromone and ethyl oleate, none of which come from the queen. NCBI Bookshelf

The hive is ruled by a king

Aristotle's king bee stood for two thousand years. Charles Butler's The Feminine Monarchie of 1609 popularised the correction that the ruling bee is female. Wikipedia, Charles Butler

All bees make honey

Only honey bees and stingless bees store honey at all. More than twenty thousand bee species are described, and the great majority store nothing a person could harvest. Discover Life

All bees live in hives

Over 90 per cent of bee species are solitary. A female digs or finds her own burrow, provisions her own cells and usually dies before her offspring emerge. There is no colony and no hive. Wikipedia, Eusociality

All bees are endangered in the same way

The western honey bee is managed livestock and is not endangered; global managed colony numbers have risen. Many wild bees genuinely are declining, and buying a hive helps the first group while competing with the second. Xerces Society

Honey is bee vomit

Nectar is drawn into the crop, a storage sac held apart from the gut by a valve, the proventriculus. It never enters the bee's stomach and is not digested, which is why a forager can carry it for hours and hand it over intact. Michigan State University

Bees are aggressive

Defensiveness is a threshold, not a temperament. African bees differ from European bees quantitatively rather than qualitatively, and their low threshold is adaptive against honey badgers, baboons, ants and honey hunters. Wikipedia, Africanized bee

A smoker calms bees by putting them to sleep

Smoke masks the alarm pheromone released by guards and injured bees, and triggers a feeding response in anticipation of fire so the bees gorge on honey. Neither is sedation, and neither lasts. Wikipedia, Bee smoker

Bees are deaf

They have no tympanal ear and cannot hear pressure waves, but they detect air particle movement through Johnston's organ in the antenna, which is exactly how the waggle dance is read in the dark. J Neuroscience, 2017

The round dance is a separate dance

There is one dance. Direction is always encoded and simply cannot be expressed well when the run is very short, so a close target produces what looks like a circle. Wikipedia, Round dance

Bees find food by smell, not by the dance

Adrian Wenner's challenge was serious and it was properly tested. Recruits carried away from the hive flew the dance vector onto empty ground where no feeder odour could exist. Riley et al., Nature, 2005

Every dance follower flies where she is told

Most do not. In one experiment 93 per cent of followers ignored the directions and returned to ground they already knew, and estimates of how often dance information is actually used sit near 10 per cent. Wikipedia, Waggle dance

The scatter in the dance is deliberate

The tuned error hypothesis predicted that advertised patch size stays constant with range. It does the opposite and grows with distance. Current reading is a sensory constraint, and the point remains contested. Frontiers, 2015

A bee flies 800 km in her lifetime

No primary measurement for that round figure could be traced. What is documented is that wing damage cuts flight capacity and shortens foraging range. Treat the number as unverified. J Insect Physiol, 2014

A 500 g jar takes 25,000 crop loads

That figure assumes nectar at 40 per cent sugar. At the concentration nectar actually arrives, the answer is 33,000 to 50,000 loads, and the flying distance figures that travel with it disagree by a factor of two. Wikipedia, Honey

Bees are born from dead oxen

Bugonia was taken as fact by Virgil and Ovid. The likeliest explanation is the drone fly Eristalis tenax, a bee mimic that breeds in carrion, plus the honey bee habit of nesting in cavities including carcasses. Wikipedia, Bugonia

Bee venom therapy treats disease

There is no scientific evidence that it treats or alters the course of cancer or any other disease, and trials in multiple sclerosis found it ineffective and capable of worsening symptoms. Frontiers in Immunology, 2024

Crystallised honey has gone off

Crystallisation is a phase change driven by the glucose to fructose ratio and by temperature, and it is arguably a marker of unprocessed honey, since heavy filtering strips the particles that seed it. Wikipedia, Honey

Varroa drinks bee blood

Fifty years of textbooks said haemolymph, and it was wrong. The mite feeds primarily on the fat body, the organ handling detoxification, immunity and overwintering reserves. Ramsey et al., PNAS, 2019

Colony collapse disorder is the name for any big bee loss

The 2006 case definition was specific: adult workers gone, no dead bees, capped brood left, stores unrobbed. Reported cases faded after about 2010, and the 2024 to 2025 United States losses do not match it. Wikipedia, Colony collapse disorder

Wax moth killed the colony

Wax moth is a symptom. Healthy colonies police their own combs, so wax moth means the colony was already weak, queenless, diseased or dead, and recording it as the cause hides the real one. National Bee Unit

Bees are obviously conscious, or obviously not

Both positions overreach. More than 500 scientists have signed a declaration stating that the evidence indicates at least a realistic possibility of conscious experience in insects, and no more than that can honestly be said. Wikipedia, New York Declaration

The oldest known bee is 100 million years old

Melittosphex burmensis from Burmese amber was described as one of the two oldest bees, but later work reclassified it as an aculeate wasp. The oldest bee fossil is Cretotrigona prisca, from the Late Cretaceous of New Jersey. Wikipedia, *Melittosphex*

11

What South African law actually requires

Two obligations catch keepers out. American foulbrood is a notifiable disease, so a suspected case is reported to the national agriculture department rather than managed quietly, and confirmation is dealt with by destruction rather than treatment, because the spores outlast anything you can do to them. Annual registration with the department is compulsory, along with marking your hives and keeping records.

The second is the demarcation line between Apis mellifera capensis in the winter rainfall Cape and Apis mellifera scutellata across the rest of the country. Bees may not be transported across it in either direction. A Cape worker can lay diploid female eggs without mating and can parasitise a scutellata colony from the inside, and moving colonies north in 1992 cost an estimated fifty thousand colonies.

The notice number, its date and the parent Act are set out in the full reference, together with a note that we could reach the industry summaries of the gazette but not the gazette itself. Confirm your own obligations with the department rather than with a website.

Hive records are yours alone. What you record in Keepa about your own hives, including disease, is readable by you and by nobody else. Health information is only ever stored when you have explicitly consented to it, because a reported foulbrood case can trigger destruction orders and damage a buyer relationship.

12

Glossary

Every technical term used above, plus the practice terms a new keeper meets in a beekeeping association meeting and is too embarrassed to ask about.

178 terms

`csd`the complementary sex determiner gene. Heterozygotes develop as females; homozygotes and hemizygotes develop as males, and diploid drone larvae are eaten by workers.
Abscondingthe entire colony abandons the nest, leaving nothing behind, in response to dearth, pests, ants, fire or repeated disturbance. Common in Apis mellifera scutellata and rare in most European stock, and not the same thing as swarming.
Acarapis woodithe tracheal mite, which enters through the first thoracic pair of spiracles and lives in the tracheae.
Afterswarma secondary swarm leaving with a virgin queen after the primary swarm has gone. Also called a cast. Far more common in African bees than in European stock.
Age polyethismthe pattern by which a worker's job changes with her age, driven by gland development and juvenile hormone. Also called temporal polyethism, and it is a default rather than a rule.
Alarm pheromonethe chemical signal that recruits defenders. Isopentyl acetate from the sting apparatus is the main one and smells of banana; 2-heptanone from the mandibular glands is weaker and also marks spent flowers.
American foulbrooda notifiable brood disease caused by the spore forming bacterium Paenibacillus larvae. The spores survive decades, so confirmed cases are dealt with by destruction rather than treatment.
Ammoplanidaea family of tiny predatory wasps, sometimes called pearl wasps, now placed as the closest living relatives of bees.
Antennal lobethe first olfactory processing centre in the brain, holding about 160 glomeruli in which odour identity is coded as a spatial pattern.
Anthophilathe clade containing all bees, nested inside the superfamily Apoidea.
Apiarya site where hives are kept. In South African usage an apiary is also commonly called a bee stand or simply a stand.
Apidaethe largest bee family, holding honey bees, stingless bees, bumblebees, orchid bees, carpenter bees and many others.
Apitoxinbee venom, dominated by melittin with phospholipase A2 as the principal allergen.
Apoideathe superfamily containing bees and the apoid wasps. Bees evolved from inside this group, not alongside it.
Arrhenotokythe normal honey bee pattern in which unfertilised eggs develop into haploid males. Contrast thelytoky.
Asynchronous muscleflight muscle that oscillates faster than nerve impulses arrive, because it is stretch activated and coupled to the resonant mechanics of the thorax.
Azimuththe sun's compass bearing on the horizon, as distinct from its height in the sky. It is the reference the waggle dance uses.
Ballingworkers clustering tightly around an intruder or an unwanted queen and shivering, killing by heat and accumulated carbon dioxide.
Basitarsusthe first and largest segment of the tarsus. On the foreleg it carries the antenna cleaner notch; on the hind leg it carries the pollen brush and the auricle.
Beardingbees hanging in a mass outside the entrance on a hot evening. It is a functional cooling response, not a symptom of a problem, and a common sight on South African summer evenings.
Bee breadpollen packed into cells with nectar and saliva and fermented by lactic acid bacteria into a stable, acidified protein store. It is what nurse bees eat to run their brood food glands.
Bee spacethe clearance of 6.4 to 9.5 mm, standardly 7 ± 2 mm, that bees leave open as a passage. A wider gap gets filled with comb, a narrower one gets propolised.
Bivouacthe temporary cluster a swarm forms, usually within 50 m of the parent hive, while scouts search for a permanent cavity.
Bluegumthe South African common name for Eucalyptus, and the source of the country's commercial workhorse honey.
Brood chamberthe box or boxes holding the brood nest, as distinct from the supers above.
Brood ester pheromonea blend of ten fatty acid esters from larval salivary glands that stimulates capping and brood rearing and inhibits worker ovaries.
Brood nestthe region of comb holding eggs, larvae and pupae, held between about 33 and 36 °C.
Buzz pollinationgripping a flower and vibrating the flight muscles to shake pollen from anthers that open only by a terminal pore. Also called sonication. Honey bees cannot do it; carpenter bees and many solitary bees can.
Buzz runthe rapid zigzag run with wings buzzing that scouts perform through a swarm cluster immediately before lift off.
Capensiseveryday South African shorthand for Apis mellifera capensis, the Cape honey bee, and by extension for the social parasitism its workers cause in scutellata colonies.
Cappingthe wax lid workers place over a ripe honey cell or a mature larva. On honey it is the colony's own certification that the contents will not ferment.
Chalkbrooda brood disease caused by the fungus Ascosphaera apis, producing hard chalky mummies that rattle and are easily removed.
Chitinthe long chain polymer of N acetylglucosamine that forms the flexible base of the insect cuticle. Rigidity comes from sclerotisation, not from chitin itself.
Colony collapse disordera specific syndrome named in 2006 and 2007, defined by rapid loss of adult workers with few dead bees, capped brood left behind, and unrobbed stores. The term is now routinely misapplied to any large loss event.
Corbiculathe pollen basket, a polished concavity on the outer face of the hind tibia fringed with hairs and holding a single long central hair that pins the pellet in place. Only females have one.
Corpora allatathe paired endocrine glands in the head that are the only source of juvenile hormone.
Cuckoo beea bee that builds no nest and gathers no pollen, laying instead in another bee's provisioned cell. More than 3,000 species are known and the habit has evolved repeatedly.
Cuticlethe layered external casing of an insect, comprising a waxy epicuticle over a hardened exocuticle and a flexible endocuticle. Because it cannot expand, an adult bee does not grow.
Deartha period with no nectar flow. In South Africa the Western Cape dearth is the hot dry January to March window and the interior dearth falls in dry autumns; dearth drives robbing and absconding.
Diastasean amylase in honey with little to do chemically, valued as a heat labile quality marker. A low diastase reading with high HMF indicates overheated or old honey.
Diutinusthe winter bee, physiologically distinct through a very high vitellogenin titre and capable of living up to 20 weeks or more.
Dorsal rim areathe narrow strip of specialised ommatidia along the top of each compound eye that reads the polarisation pattern of the sky.
Drawn combfoundation that the bees have built out into finished cells. It is a valuable asset because building it costs the colony 6.6 to 10 kg of honey per kilogram of wax.
Dronethe male bee, haploid, fatherless, stingless, unable to forage or feed brood, and built for one mating flight.
Drone congregation areaan aerial lekking site 10 to 40 m up where drones from many colonies gather and virgin queens fly to mate. The same sites are used year after year although no drone survives from one season to the next.
Dysenterysoiling of the comb and hive front by bees unable to void outside. It is a symptom, not a diagnosis, and can accompany nosemosis.
Ecdysisthe shedding of the old cuticle at the end of a moult.
Electroreceptiondetection of electric fields. Bees carry a positive charge in flight and flowers a lower potential; bumblebee mechanosensory hairs and honey bee antennae have both been shown to respond.
Emergency queen cella queen cell drawn out from an ordinary worker cell after the queen is suddenly lost. Because the larva was already fed as a worker, the result is usually smaller and less prolific.
Endophallusthe drone's mating organ, everted explosively under haemolymph pressure and torn away to remain in the queen as the mating sign.
Ephemeristhe relationship between the time of day and the sun's position. Bees learn the local ephemeris by observation and apply it in landscapes they have never flown.
Ethyl oleatea primer pheromone made in the honey crop and carried by foragers, delaying young bees from starting to forage.
European foulbrooda brood disease caused by Melissococcus plutonius, attacking young unsealed larvae, watery rather than ropy, and often clearing when forage improves.
Eusocialitycooperative brood care, overlapping generations and reproductive division of labour occurring together. Over 90 per cent of bee species are solitary; eusociality is the exception.
Extrafloral nectarya nectar gland outside the flower, usually on a leaf or petiole, which buys defence by recruiting ants rather than buying pollination.
Fanningwing beating to drive a directional current of air through the nest, used to ripen honey and to cool the brood nest.
Fat bodythe diffuse tissue lining the abdominal wall that acts as liver, fat store and endocrine organ, and produces vitellogenin. It is what Varroa actually feeds on.
Flagellumthe whip like distal part of the antenna, made of flagellomeres: 10 in the female castes, 11 in the drone.
Flicker fusionthe frequency at which a flickering light appears steady. Bees reach 200 Hz at 3000 lux against about 60 Hz in humans, which is what makes optic flow odometry possible.
Floral constancythe tendency of an individual forager to work one flower species while it remains profitable.
Flowa period when nectar is coming in strongly. South African beekeepers speak of the eucalyptus flow, the fynbos flow and the sunflower flow as distinct events.
Footprint pheromonean oily secretion deposited from the tarsal glands as the bee walks. The queen's version inhibits queen cup building, and its decline signals a failing queen.
Foundationa sheet of wax or plastic embossed with cell bases, hung in a frame for the bees to draw out. Commercial worker foundation runs 5.20 to 5.75 mm against 5.03 to 5.55 mm in wild comb.
Fynbosthe fire adapted shrubland of the Cape Floristic Region, source of the flagship Cape honey and the natural range of Apis mellifera capensis.
Glossathe tongue, hairy and tipped with a spoon shaped labellum, which laps and draws nectar.
Glucose oxidasethe enzyme that converts glucose into gluconic acid and hydrogen peroxide. It is largely inactive in ripe honey and switches on when honey is diluted, exactly when a microbial threat becomes real.
Guard beea worker in the short phase just before foraging who antennates every incoming bee at the entrance. Five to ten are typically on duty at once.
Haemolymphinsect blood, clear to straw coloured, circulating in an open body cavity and carrying nutrients, hormones and immune cells but no oxygen.
Hamulithe row of hooks on the leading edge of the hind wing that catch into a fold on the forewing, locking the two into a single aerofoil in flight.
Haplodiploidythe genetic system in which males develop from unfertilised haploid eggs and females from fertilised diploid ones. Its role in the evolution of eusociality is now seriously contested.
Harmonic radara tracking method in which a bee carries a passive transponder that reradiates at twice the transmit frequency, so it stands out against ground clutter that does not answer on the harmonic.
Heater beea worker who shivers her flight muscles without moving her wings, pressing her thorax against sealed cappings or climbing into empty cells inside the brood slab to heat from within.
HMFhydroxymethylfurfural, formed as fructose degrades in acid. It accumulates with heat and time, so it is a combined index of thermal abuse and age.
Honeycomb conjecturethe proposition, first recorded by Varro in 36 BC and proved by Hales in 1999, that among equal area shapes tiling a plane the regular hexagon minimises total wall material.
Honeydewthe sugar surplus excreted by aphids and other sap feeders, worked by bees into a dark, mineral rich honey that Codex specifies separately.
Hybrid zonethe 100 to 200 km band between about 31°S and 33°S where capensis and scutellata meet. Traits vary clinally across it and colony movement across it is regulated.
Hygienic behaviourthe detection and removal of diseased or infested brood by workers. It is one mechanism behind the mite resistance of savannah honey bees.
Hypopharyngeal glandthe paired head gland that produces brood food in a young worker and the enzyme invertase in a forager.
Instara stage between larval moults. A worker larva passes through five.
Invertasethe enzyme, also called sucrase, that splits sucrose into glucose and fructose, roughly doubling the number of dissolved particles and so sharply raising osmotic pressure.
Isopentyl acetatethe principal alarm pheromone, released from the sting apparatus, smelling of banana and marking the target for further stings.
Johnston's organthe chordotonal organ in the antennal pedicel that detects deflection of the flagellum. It is how a bee reads the waggle dance in the dark, and it also registers gravity, airflow and electric field changes.
Juvenile hormonethe hormone from the corpora allata that drives caste differentiation and the shift from nurse to forager. It is mutually antagonistic with vitellogenin.
Kenyon cellthe densely packed intrinsic neuron of the mushroom body. A honey bee has about 170,000 per mushroom body against 2,500 in a fruit fly.
Langstroth hivethe movable frame hive that applies bee space around every surface of a hanging frame, patented in 1852 and the basis of essentially all modern hive management.
Laying workera worker whose ovaries have activated in the absence of a queen and open brood. The signature is multiple eggs per cell, eggs off centre on the cell wall, patchy brood and domed drone cappings in worker cells.
Leading edge vortexthe attached vortex that forms along the front of a flapping wing and holds flow attached at angles that would stall a fixed wing. It is the main reason bees fly.
Malpighian tubuleone of the 100 to 150 fine tubes floating in the haemolymph that convert nitrogenous waste to uric acid, which costs almost no water to excrete.
Mandibular glandthe gland at the base of each mandible. It makes royal jelly lipids in a young worker, 2-heptanone in a forager, and queen mandibular pheromone in a queen.
Mating signthe ruptured endophallus left in the queen after a drone mates and dies.
Meconiumthe waste accumulated in a larva's closed midgut through the whole of larval life and voided into the base of the cell only after capping. Successive deposits are what darken old brood comb.
Meliponinithe tribe of stingless bees, between 462 and 552 described species, of which Africa holds roughly 36. They store food in separate pots rather than comb.
Migratory beekeepingmoving colonies between sites to follow flows or fulfil pollination contracts. In South Africa it is also how the Cape bee parasite reached scutellata country.
Mushroom bodiesthe paired brain structures that are the seat of olfactory learning and of short and long term memory.
Nasonov glandthe scent gland between the last two tergites, exposed by flexing the abdomen down and fanning. Its lemony, citral dominated scent marks the entrance, a settled swarm or a water source.
Nectarthe sugar rich liquid plants secrete to buy pollination, arriving at the hive at 70 to 80 per cent water.
Nectar guidea pattern on a petal, often in ultraviolet absorbing pigment invisible to humans, that functions as a landing target.
Neonicotinoida systemic insecticide acting as an agonist at the insect nicotinic acetylcholine receptor. Being systemic and water soluble, it reaches nectar and pollen by design rather than by accident.
Nosemosisan adult gut disease caused by microsporidians of the genus Nosema, moved by a contested 2020 revision into Vairimorpha, so both names are current. Diagnosis needs microscopy.
Nuca small colony, usually on three to five frames, used to start a new hive, hold a mated queen or make an increase. Short for nucleus colony.
Nurse beea worker of roughly days 3 to 11 whose hypopharyngeal glands are at maximum output and who feeds the brood.
Ocellusone of the three simple eyes on the crown. Each has a single lens, forms no image, and reports light intensity and horizon for flight stabilisation.
Ommatidiumone facet unit of the compound eye, sampling a narrow cone of the visual field. A worker has roughly 4,000 to 6,000 per eye.
Optic flowthe image motion sweeping across the eye during flight. Bees measure distance by integrating it, which is why a flight over featureless water is underreported by a factor of three to four.
Orientation flighta deliberate learning flight taken by a young bee before she forages, building the landmark memories she will navigate by.
Ovarioleone of the parallel tubes making up an ovary, in which eggs mature in series. A queen has 120 to 200 per ovary against 2 to 12 in a worker.
Ovipositorthe egg laying organ of a female insect. The sting is a modified ovipositor, which is why no drone has one.
Paenibacillus larvaethe spore forming bacterium that causes American foulbrood.
Path integrationcontinuously combining heading and distance travelled into a running estimate of the straight line home vector. Also called dead reckoning.
Peritrophic membranethe chitin and protein sheath surrounding the food bolus in the midgut, protecting the epithelium from abrasion by pollen. It is the gut's soft point and the entry route for Nosema.
Pheromonea chemical signal between individuals of one species. A honey bee colony runs on roughly twenty such channels, from queen mandibular pheromone to brood ester pheromone.
Pipinga vibratory signal. A free virgin queen toots, a queen still sealed in her cell quacks in reply, and a swarm scout pipes at nestmates to warm their flight muscles before departure.
Plumose setaa branched, feathery body hair. The branching multiplies surface area, traps pollen electrostatically and can register a flower's electric field.
Polarised lightskylight whose electric field vectors are arranged tangentially around the sun. Reading its orientation in one patch of blue sky reveals where the sun is, even when the sun is hidden.
Pollen pressthe rastellum and auricle acting together to squeeze pollen upward and outward into the corbicula as the bee flexes her hind leg.
Prepupathe stage after a larva uncurls, stretches lengthwise, spins its cocoon and defecates for the first time, before the pupal form appears.
Proboscisthe temporary sucking tube formed on demand when the galeae and labial palps close around the glossa. It folds back under the head when feeding ends.
Proboscis extension reflexthe reflexive extension of the proboscis when sugar touches the antennae. Pairing an odour with the sugar turns the reflex into a readable laboratory measure of learning.
Propodeumthe first true abdominal segment, fused onto the back of the thorax in Hymenoptera, which is why the visible thorax and the visible abdomen do not match the true segment counts.
Propolisbee glue, made from plant resin mixed with wax and enzymes. It seals gaps of about 6 mm or less, varnishes surfaces, and mummifies carcasses too heavy to remove.
Proventriculusthe valve between crop and midgut, carrying a toothed structure whose four lips comb pollen out of the nectar in the crop. It is why a forager can carry nectar for hours undigested.
Pseudoqueena laying worker producing queen like mandibular secretions and winning reproductive competition inside a colony. In Apis mellifera capensis the pseudoqueen's daughters are clonal copies doing the same.
Pupathe sealed, immobile stage in which adult structures develop from imaginal discs. In a worker it lasts 12 days.
Queen cella vertical, peanut shaped cell about 25 mm long. Swarm cells hang from the comb margins, supersedure cells from the comb face, emergency cells are drawn out from ordinary worker cells.
Queen cupan empty queen cell base, built along the comb margins year round. It means nothing until the queen lays in it.
Queen excludera grid sized to pass workers but not the queen, used to keep brood out of the honey supers. Removing them from experimental hives in Brazil in 1957 is how 26 scutellata swarms escaped.
Queen mandibular pheromonethe queen's principal social signal, with five major components including 9-ODA. It suppresses queen rearing and worker ovaries, forms the retinue and holds a swarm together.
Queenrighthaving a functioning laying queen. The chapters make the point that this is a chemical state with a gradient rather than a binary.
Quorum sensingreaching a decision when a threshold number of individuals is present rather than by consensus. A swarm commits when 20 to 30 scouts are at one site simultaneously.
Rastellumthe comb of stiff spines along the lower edge of the hind tibia that rakes pollen from the opposite leg's brush.
Requeeningreplacing a colony's queen with a new one. North of the hybrid zone, laying workers producing worker brood in a queenless colony are a parasite incident, not a requeening problem.
Retinuethe ring of about 17 to 18 workers surrounding the queen at any moment, licking, grooming and feeding her and carrying her pheromone away on their bodies.
Robbingcolonies raiding another colony's stores, typically in a dearth. Guards tighten their acceptance threshold rapidly when robbing pressure rises.
Round dancethe tight alternating circles a forager performs for very close food. It is now understood as the short distance end of one continuous dance rather than a separate dance.
Royal jellythe glandular secretion of nurse bees, about 67 per cent water and 12.5 per cent protein, fed to all young larvae and to a queen larva without interruption for her whole larval life.
Royalactinthe name Kamakura gave in 2011 to monomeric MRJP1, proposed as sufficient to induce queen differentiation. Rebutted in 2016 in the same journal and still contested.
Sacbrooda viral brood disease in which the larva becomes a fluid filled sac that lifts out intact, with a darker upturned head and no odour.
Scleritea hardened plate of the exoskeleton. A tergite is dorsal, a sternite ventral, and flexible membrane between them lets the abdomen telescope.
Sclerotisationthe tanning process that embeds chitin in a hardened protein matrix. A soft white pupa and a hard brown forager are the same material at different degrees of it.
Scopaa brush of pollen carrying hairs, on the legs in most families and under the abdomen in Megachilidae. It is a brush, not a basket, and carries pollen dry.
Scoutan experienced forager searching for a new nest site or a new forage patch. Twenty to fifty scouts serve a swarm.
Scutellataeveryday South African shorthand for Apis mellifera scutellata, the savannah bee of the summer rainfall interior and the whole country outside the Cape.
Shaking signala worker grasping a nestmate and vibrating at about 16 Hz for one or two seconds, raising general activity. Also called dorsoventral abdominal vibration.
Shimmeringthe defensive wave of hundreds of Apis dorsata flipping their abdomens upward in under 400 ms, which keeps hovering hornets about 50 cm away.
Shot brooda scattered pepperpot pattern of empty cells in an otherwise solid brood slab, caused by workers eating diploid drone larvae. It is a readable signal of inbreeding.
Skepa straw basket hive with comb fixed to its walls, so the comb cannot be inspected or replaced and harvesting usually killed the colony. Superseded by the movable frame.
Small hive beetleAethina tumida, native to sub Saharan Africa and a minor scavenger here, but destructive where it has invaded. The damage is fermentation by a yeast the larvae carry, not chewing.
Smokerthe tool that produces cool smoke to calm a colony during inspection. Smoke is also a documented absconding trigger, so restraint matters in a dearth.
Snapshot memorya stored retinotopic view of the panorama as seen from a goal, which the bee reduces the mismatch against as she approaches.
Spermathecathe spherical sac about 1 mm across in which a queen stores the sperm of her whole life. Opaque and white in a laying queen, clear in an unmated one.
Spiracleone of the ten valved pairs of openings through which air enters the tracheal system.
Sting extension reflexthe aversive counterpart to the proboscis extension reflex, conditioned by pairing an odour with a mild shock. A bee can hold appetitive and aversive memories for two odours at once.
Stingless beea meliponine, with a vestigial sting, defending by biting. Their honey is markedly wetter at 25 to 35 per cent water and ferments quickly, so it is kept cool and eaten fresh.
Stop signala brief buzz of about 150 ms at 350 to 450 Hz delivered as a head butt, which makes a dancer quit the floor. It suppresses recruitment to crowded, depleted or dangerous sources and breaks deadlock in swarm decisions.
Strigilisthe antenna cleaner on the foreleg, a semicircular notch in the basitarsus closed by a tibial spur. A blocked antenna is a blind bee.
Subesophageal ganglionthe ganglion below the oesophagus controlling mouthparts and salivary glands.
Subgenual organthe vibration receptor in the leg, which detects signals carried through the comb.
Supera box of comb placed above the brood chamber for honey storage. The verb, to super, means to add one.
Supersedurethe colony replacing a failing queen without swarming. The old queen is not usually killed at once, and mother and daughter often lay side by side for weeks.
Swarmthe reproductive division of a colony, the old queen leaving with 60 to 75 per cent of the workers. In South African usage the word is also used loosely for a colony itself, as in the national estimate of about 200,000 managed swarms.
Swarm cella queen cell built along the bottom and side margins of the comb when a colony is preparing to reproduce, usually three or more and of varying age.
Tergitethe dorsal plate of a body segment, overlapping the one behind it.
Thelytokyparthenogenesis in which unfertilised eggs develop into diploid females. In Apis mellifera capensis it is automictic with central fusion, and it is the root of the social parasitism problem.
Thermoregulationthe colony's control of nest temperature, by shivering to heat and by fanning, water spreading and bearding to cool. A brood nest can be held near 35 °C with ambient well above 40 °C.
Time memorythe trained recall of when a food source was available. Bees hold about five or six such memories routinely, and the same clock compensates the sun compass.
Tracheaone of the cuticle lined air tubes, held open by spiral thickenings, that branch into tracheoles ending directly on the tissues. Gas exchange never enters the blood.
Tremble dancea slow shuddering walk performed by a forager who cannot find a receiver. It recruits more receivers and suppresses further foraging: the loading bay is full.
Trophallaxismouth to mouth food transfer. It is better understood as the colony's data bus than as feeding, since it also moves floral scent, hormones and chemical cues.
Tuned error hypothesisthe proposal that scatter in the waggle dance is deliberate, so a patch of constant size is advertised at any range. It has not held up, since advertised patch size increases with distance.
Undertakingthe removal of dead bees from the nest, a task of workers around days 12 to 17. A carcass too heavy to remove is encased in propolis instead.
Varroa destructorthe obligate ectoparasitic mite that jumped from Apis cerana to Apis mellifera in the twentieth century. It feeds primarily on the fat body, not on haemolymph, and vectors and amplifies viruses.
Vitellogeninthe glycolipoprotein made in the fat body that acts as food store, antioxidant and behavioural hormone. Very high titres are what make a winter bee live months rather than weeks.
Waggle dancethe figure of eight run on vertical comb encoding direction as an angle from vertical measured against gravity, and distance as the duration of the straight waggle phase.
Wax mirrorthe smooth translucent oval on a sternite through which liquid wax is secreted, setting outside as a scale.
Wax mothGalleria mellonella and Achroia grisella, whose larvae tunnel through comb under silk galleries. Wax moth is a symptom of a weak or dead colony, not a cause of its loss.
Winter clusterthe two layer mass a colony forms below roughly 10 to 14 °C ambient, with a dense mantle and a looser heat generating core. Recent work argues the mantle is thermally stressed rather than a warm blanket.
Worker policingworkers eating eggs laid by other workers, which is what keeps worker reproduction rare in a queenright colony.
Xylocopathe carpenter bees, which excavate galleries in dead sound wood, are powerful buzz pollinators and, lacking bumblebees, are South Africa's obvious candidate for managed buzz pollination.

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Where this came from

This page is drawn from a reference document held with the project, running to about a hundred thousand words across eleven chapters, compiled from published encyclopedias, university extension services, regulators and peer reviewed papers, then checked line by line against independent sources. Roughly fifty claims were corrected or removed at that stage, including several that are widely repeated.

Two rules were kept throughout. Where sources disagree, both figures are given and the disagreement is named. Where a figure could not be confirmed against a primary source, it says so in the sentence. The national colony count is the clearest example: South Africa has not run a census since 1975, so any number you see quoted for it, including ours, is an estimate standing on nothing.

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