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.
`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.