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105-500: They traditionally lived in small groups, farming for three to five years, then moving on as the fertility of the soil diminished (Stauder 1971). They were active bee keepers, collecting honey from hives consisting of hollowed logs placed in trees. They did some hunting and snaring of game and trapping of fish. They raised the bulk of their own food by farming, animals providing only a small part of their diet. Food production has changed since Stauder's time. The single most obvious change

210-401: A division of labour into reproductive and non-reproductive adults, plus overlapping generations. This division of labour creates specialized groups within eusocial societies which are called castes . In some species, groups of cohabiting females may be sisters, and if there is a division of labour within the group, they are considered semisocial . The group is called eusocial if, in addition,

315-464: A glassy state and becomes an amorphous solid (noncrystalline). The viscosity of honey is affected greatly by both temperature and water content. The higher the water percentage, the more easily honey flows . Above its melting point, however, water has little effect on viscosity. Aside from water content, the composition of most types of honey also has little effect on viscosity. At 25 °C (77 °F), honey with 14% water content generally has

420-598: A supercooled solution of various sugars in water. These concentrations of sugar can only be achieved near room temperature by evaporation of a less concentrated solution, in this case nectar. For osmotic reasons such high concentrations of sugar are extremely unfavorable to microbiological reproduction and all fermentation is consequently halted. The bees then cap the cells of finished honey with wax. This seals them from contamination and prevents further evaporation. So long as its water concentration does not rise much above 18%, honey has an indefinite shelf life, both within

525-663: A bee, be it a solitary or social species, involves the laying of an egg, the development through several moults of a legless larva , a pupation stage during which the insect undergoes complete metamorphosis , followed by the emergence of a winged adult. The number of eggs laid by a female during her lifetime can vary from eight or less in some solitary bees, to more than a million in highly social species. Most solitary bees and bumble bees in temperate climates overwinter as adults or pupae and emerge in spring when increasing numbers of flowering plants come into bloom. The males usually emerge first and search for females with which to mate. Like

630-429: A compartment (a "cell") with an egg and some provisions for the resulting larva, then seals it off. A nest may consist of numerous cells. When the nest is in wood, usually the last (those closer to the entrance) contain eggs that will become males. The adult does not provide care for the brood once the egg is laid, and usually dies after making one or more nests. The males typically emerge first and are ready for mating when

735-466: A distinct set of genera which are commonly known by their nesting behavior or preferences, namely: carpenter bees, sweat bees , mason bees, plasterer bees , squash bees , dwarf carpenter bees , leafcutter bees, alkali bees and digger bees . Most solitary bees are fossorial , digging nests in the ground in a variety of soil textures and conditions, while others create nests in hollow reeds or twigs, or holes in wood . The female typically creates

840-408: A few types have non-Newtonian viscous properties. Honeys from heather or manuka display thixotropic properties. These types of honey enter a gel-like state when motionless, but liquefy when stirred. Because honey contains electrolytes , in the form of acids and minerals, it exhibits varying degrees of electrical conductivity . Measurements of the electrical conductivity are used to determine

945-584: A five-string lyre , thumb piano , drum, rattles, panflute . In addition, they play a vertically suspended marimba with as few as three wooden bars, but this is seen as a way of passing time, especially when guarding fields, rather than an instrument for music. Their vocal music includes singing of both harmonies and antiphonal parts. Often, this results in two parts being sung by women and two parts by men. Changes are happening rapidly to their traditional way of life. Since about 1971, many Majangir have become Christians (Hoekstra 2003 and Sato 2002). Further, since

1050-555: A food for their larvae . To this end bees stockpile honey to provide for themselves during ordinary foraging as well as during lean periods, as in overwintering . During foraging bees use part of the nectar they collect to power their flight muscles. The majority of nectar collected is not used to directly nourish the insects but is instead destined for regurgitation , enzymatic digestion , and finally long-term storage as honey. During cold weather or when other food sources are scarce, adult and larval bees consume stored honey, which

1155-469: A honey, which may be described as "sweet", "flowery", "citrus", "almond" or "rancid", among other terms. In addition, VOCs play a large role in determining the specific flavor of the honey, both through the aromas and flavor. VOCs from honeys in different geographic regions can be used as floral markers of those regions, and as markers of the bees that foraged the nectars. Honey is classified by its source (floral or not), and divisions are made according to

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1260-525: A larva's food is supplied gradually as it develops, as is the case in honey bees and some bumblebees. Most other bees, including familiar insects such as carpenter bees , leafcutter bees and mason bees are solitary in the sense that every female is fertile, and typically inhabits a nest she constructs herself. There is no division of labor so these nests lack queens and worker bees for these species. Solitary bees typically produce neither honey nor beeswax . Bees collect pollen to feed their young, and have

1365-440: A length of 39 millimetres (1.54 in). Bees feed on nectar and pollen , the former primarily as an energy source and the latter primarily for protein and other nutrients. Most pollen is used as food for their larvae . Vertebrate predators of bees include primates and birds such as bee-eaters ; insect predators include beewolves and dragonflies . Bee pollination is important both ecologically and commercially , and

1470-695: A long and varied history, with its beginnings in prehistoric times . Several cave paintings in Cuevas de la Araña in Spain depict humans foraging for honey at least 8,000 years ago. While Apis mellifera is an Old World insect , large-scale meliponiculture of New World stingless bees has been practiced by Mayans since pre-Columbian times. Honey is produced by bees who have collected nectar or honeydew . Bees value honey for its sugars, which they consume to support general metabolic activity , especially that of their flight muscles during foraging , and as

1575-729: A male is haploid (has only one copy of each gene), his daughters (which are diploid , with two copies of each gene) share 100% of his genes and 50% of their mother's. Therefore, they share 75% of their genes with each other. This mechanism of sex determination gives rise to what W. D. Hamilton termed "supersisters", more closely related to their sisters than they would be to their own offspring. Workers often do not reproduce, but they can pass on more of their genes by helping to raise their sisters (as queens) than they would by having their own offspring (each of which would only have 50% of their genes), assuming they would produce similar numbers. This unusual situation has been proposed as an explanation of

1680-436: A possible fire), making them less aggressive, and obscures the pheromones the bees use to communicate. The honeycomb is removed from the hive and the honey may be extracted from it either by crushing or by using a honey extractor . The honey is then usually filtered to remove beeswax and other debris. Before the invention of removable frames, bee colonies were often sacrificed to conduct the harvest. The harvester would take all

1785-631: A primary role in determining honey flavors and aromas . VOCs are carbon-based compounds that readily vaporize into the air, providing aroma, including the scents of flowers, essential oils, or ripening fruit. The typical chemical families of VOCs found in honey include hydrocarbons , aldehydes , alcohols , ketones , esters , acids , benzenes , furans , pyrans , norisoprenoids , and terpenes , among many others and their derivatives. The specific VOCs and their amounts vary considerably between different types of honey obtained by bees foraging on different plant sources. By example, when comparing

1890-449: A primary role in determining the exact temperature. Of these acids, the amino acids, which occur in very small amounts, play an important role in the darkening of honey. The amino acids form darkened compounds called melanoidins , during a Maillard reaction . The Maillard reaction occurs slowly at room temperature, taking from a few to several months to show visible darkening, but speeds up dramatically with increasing temperatures. However,

1995-533: A single season colony cycle, even in the tropics, and only mated females hibernate. A few species have long active seasons and attain colony sizes in the hundreds, such as Halictus hesperus . Some species are eusocial in parts of their range and solitary in others, or have a mix of eusocial and solitary nests in the same population. The orchid bees (Apidae) include some primitively eusocial species with similar biology. Some allodapine bees (Apidae) form primitively eusocial colonies, with progressive provisioning :

2100-518: A solid), so melting crystallized honey can easily result in localized caramelization if the heat source is too hot or not evenly distributed. However, honey takes substantially longer to liquefy when just above the melting point than at elevated temperatures. Melting 20 kg (44 lb) of crystallized honey at 40 °C (104 °F) can take up to 24 hours, while 50 kg (110 lb) may take twice as long. These times can be cut nearly in half by heating at 50 °C (122 °F); however, many of

2205-418: A solution of fructose and other ingredients. The density of honey typically ranges between 1.38 and 1.45 kg/L at 20 °C. The melting point of crystallized honey is between 40 and 50 °C (104 and 122 °F), depending on its composition. Below this temperature, honey can be either in a metastable state, meaning that it will not crystallize until a seed crystal is added, or, more often, it

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2310-463: A viscosity around 400  poise , while a honey containing 20% water has a viscosity around 20 poise. Viscosity increases very slowly with moderate cooling; a honey containing 16% water, at 70 °C (158 °F), has a viscosity around 2 poise, while at 30 °C (86 °F), the viscosity is around 70 poise. With further cooling, the increase in viscosity is more rapid, reaching 600 poise at around 14 °C (57 °F). However, while honey

2415-512: Is Cretotrigona prisca , a corbiculate bee of Late Cretaceous age (~70 mya) found in New Jersey amber . A fossil from the early Cretaceous (~100 mya), Melittosphex burmensis , was initially considered "an extinct lineage of pollen-collecting Apoidea sister to the modern bees", but subsequent research has rejected the claim that Melittosphex is a bee, or even a member of the superfamily Apoidea to which bees belong, instead treating

2520-471: Is a sweet and viscous substance made by several species of bees , the best-known of which are honey bees . Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primarily floral nectar ) or the secretions of other insects, like the honeydew of aphids. This refinement takes place both within individual bees, through regurgitation and enzymatic activity, and during storage in

2625-414: Is also affected by water content, because a high percentage of water inhibits crystallization, as does a high dextrin content. Temperature also affects the rate of crystallization, with the fastest growth occurring between 13 and 17 °C (55 and 63 °F). Crystal nuclei (seeds) tend to form more readily if the honey is disturbed, by stirring, shaking, or agitating, rather than if left at rest. However,

2730-607: Is an important food for virtually all hunter-gatherer cultures in warm climates, with the Hadza people ranking honey as their favorite food. Honey hunters in Africa have a mutualistic relationship with certain species of honeyguide birds. Possibly the world's oldest fermented beverage , dating from 9,000 years ago, mead ("honey wine") is the alcoholic product made by adding yeast to honey-water must and fermenting it for weeks or months. The yeast Saccharomyces cerevisiae

2835-476: Is believed that mites eat fungi that attack pollen, so the relationship in this case may be mutualistic . Molecular phylogeny was used by Debevic et al , 2012, to demonstrate that the bees ( Anthophila ) arose from deep within the Crabronidae sensu lato , which was thus rendered paraphyletic . In their study, the placement of the monogeneric Heterogynaidae was uncertain. The small family Mellinidae

2940-408: Is collected from wild bee colonies or from domesticated beehives. On average, a hive will produce about 29 kilograms (65 lb) of honey per year. Wild bee nests are sometimes located by following a honeyguide bird. To safely collect honey from a hive, beekeepers typically pacify the bees using a bee smoker . The smoke triggers a feeding instinct (an attempt to save the resources of the hive from

3045-570: Is commonly used in modern mead production. Mead varieties include drinks called metheglin (with spices or herbs), melomel (with fruit juices, such as grape, specifically called pyment ), hippocras (with cinnamon ), and sack mead (high concentration of honey), many of which have been developed as commercial products numbering in the hundreds in the United States. Honey is also used to make mead beer , called "braggot". The physical properties of honey vary, depending on water content,

3150-594: Is in a "labile" state, being saturated with enough sugars to crystallize spontaneously. The rate of crystallization is affected by many factors, but the primary factor is the ratio of the main sugars: fructose to glucose. Honeys that are supersaturated with a very high percentage of glucose, such as brassica honey, crystallize almost immediately after harvesting, while honeys with a low percentage of glucose, such as chestnut or tupelo honey, do not crystallize. Some types of honey may produce few but very large crystals, while others produce many small crystals. Crystallization

3255-415: Is in their own honey stomachs, the hive bees regurgitate the nectar, repeatedly forming bubbles between their mandibles , speeding its digestion and concentration. These bubbles create a large surface area per volume and by this means the bees evaporate a portion of the nectar's water into the warm air of the hive. Hive bees form honey-processing groups. These groups work in relay, with one bee subjecting

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3360-605: Is limiting access to humidity. In its cured state, honey has a sufficiently high sugar content to inhibit fermentation. If exposed to moist air, its hydrophilic properties pull moisture into the honey, eventually diluting it to the point that fermentation can begin. The long shelf life of honey is attributed to an enzyme found in the stomach of bees. The bees mix glucose oxidase with expelled nectar they previously consumed, creating two byproducts – gluconic acid and hydrogen peroxide , which are partially responsible for honey acidity and suppression of bacterial growth. Honey

3465-399: Is many times as energy-dense as the nectar from which it is made. After leaving the hive, a foraging bee collects sugar-rich nectar or honeydew. Nectar from the flower generally has a water content of 70 to 80% and is much less viscous than finished honey, which usually has a water content around 18%. The water content of honeydew from aphids and other true bugs is generally very close to

3570-623: Is minor and does not seriously impede communication The language has implosive consonants (bilabial and retroflex), but no ejective consonants (Bender 1983). There are seven vowels, and length of duration of the vowel is also distinctive, such as goopan 'punishment' and gopan 'road'. In addition, two tones also distinguish meaning, on both the word level and the grammatical level: táŋ (higher tone) 'cow', tàŋ (lower tone) 'abscess'. Bee Apiformes (from Latin 'apis' ) Bees are winged insects closely related to wasps and ants , known for their roles in pollination and, in

3675-421: Is normal. At very low temperatures, honey does not freeze solid; rather its viscosity increases. Like most viscous liquids , the honey becomes thick and sluggish with decreasing temperature. At −20 °C (−4 °F), honey may appear or even feel solid, but it continues to flow at very low rates. Honey has a glass transition between −42 and −51 °C (−44 and −60 °F). Below this temperature, honey enters

3780-573: Is often pasteurized , to kill any yeast, by heating it above 70 °C (158 °F). Like all sugar compounds, honey caramelizes if heated sufficiently, becoming darker in color, and eventually burns. However, honey contains fructose, which caramelizes at lower temperatures than glucose. The temperature at which caramelization begins varies, depending on the composition, but is typically between 70 and 110 °C (158 and 230 °F). Honey also contains acids, which act as catalysts for caramelization. The specific types of acids and their amounts play

3885-598: Is preferred because all bee colonies, even those in very large agricultural apiculture operations, readily leave the protection of humans in swarms that can establish successful wild colonies. Much of the effort in commercial beekeeping is dedicated to persuading a hive that is ready to swarm to produce more honeycomb in its present location. This is usually done by adding more space to the colony with honey supers , empty boxes placed on top of an existing colony. The bees can then usually be enticed to develop this empty space instead of dividing their colony through swarming. Honey

3990-441: Is sometimes adulterated by the addition of other sugars, syrups, or compounds to change its flavor or viscosity, reduce cost, or increase the fructose content to inhibit crystallization . Honey has been adulterated since ancient times, when honey was sometimes blended with plant syrups such as maple , birch , or sorghum and sold to customers as pure honey. Sometimes crystallized honey was mixed with flour or other fillers, hiding

4095-605: Is sweet because of its high concentrations of the monosaccharides fructose and glucose . It has about the same relative sweetness as sucrose (table sugar). One standard tablespoon (15 mL) of honey provides around 190 kilojoules (46 kilocalories) of food energy . It has attractive chemical properties for baking and a distinctive flavor when used as a sweetener. Most microorganisms cannot grow in honey and sealed honey therefore does not spoil . Samples of honey discovered in archaeological contexts have proven edible even after millennia. Honey use and production has

4200-461: Is that people are now living in permanent settlements. Livestock was not traditionally raised, but many Majangir have begun small scale livestock raising since about 1980. In addition, they have begun planting fruit and coffee trees, plants that take a number of years to produce a crop. The Majangir traditionally had a very egalitarian society, with no standing political leaders (Stauder 1971, 1972). The only people in official positions were people in

4305-444: Is the ancestral state for all eusocial species so far investigated, so it is likely that haplodiploidy contributed to the evolution of eusociality in bees. Bees may be solitary or may live in various types of communities. Eusociality appears to have originated from at least three independent origins in halictid bees. The most advanced of these are species with eusocial colonies; these are characterised by cooperative brood care and

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4410-555: Is thought to have occurred due to rapid climatic cooling around the Eocene-Oligocene boundary , leading to the extinction of some bee lineages such as the tribe Melikertini . Over the Paleogene and Neogene , different bee lineages continued to spread all over the world, and the shifting habitats and connectedness of continents led to the isolation and evolution of many new bee tribes. The oldest non-compression bee fossil

4515-399: Is viscous, it has low surface tension of 50–60 mJ/m , making its wettability similar to water, glycerin , or most other liquids. The high viscosity and wettability of honey cause stickiness , which is a time-dependent process in supercooled liquids between the glass-transition temperature (T g ) and the crystalline-melting temperature. Most types of honey are Newtonian liquids , but

4620-541: The Early Cretaceous (about 124 million years ago) on the supercontinent of West Gondwana , just prior to its breakup into South America and Africa . The supercontinent is thought to have been a largely xeric environment at this time; modern bee diversity hotspots are also in xeric and seasonal temperate environments, suggesting strong niche conservatism among bees ever since their origins. Genomic analysis indicates that despite only appearing much later in

4725-707: The Stenotritidae ), and by the end of the Cretaceous, South American bees had also colonized North America. The North American fossil taxon Cretotrigona belongs to a group that is no longer found in North America, suggesting that many bee lineages went extinct during the Cretaceous-Paleogene extinction event . Following the K-Pg extinction, surviving bee lineages continued to spread into

4830-443: The nucleation of microscopic seed-crystals is greatest between 5 and 8 °C (41 and 46 °F). Therefore, larger but fewer crystals tend to form at higher temperatures, while smaller but more-numerous crystals usually form at lower temperatures. Below 5 °C, the honey will not crystallize, thus the original texture and flavor can be preserved indefinitely. Honey is a supercooled liquid when stored below its melting point, as

4935-588: The Bumblebee Specialist Group to review the threat status of all bumblebee species worldwide using the IUCN Red List criteria. There are many more species of primitively eusocial than highly eusocial bees, but they have been studied less often. Most are in the family Halictidae , or "sweat bees". Colonies are typically small, with a dozen or fewer workers, on average. Queens and workers differ only in size, if at all. Most species have

5040-597: The Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale. The Halictidae first appear in the Early Eocene with species found in amber. The Stenotritidae are known from fossil brood cells of Pleistocene age. The earliest animal-pollinated flowers were shallow, cup-shaped blooms pollinated by insects such as beetles , so the syndrome of insect pollination was well established before

5145-543: The Majang area. "This has led to the pauperization of the Majang and the looming environmental disaster of the massive deforestation of the Majang forest, part of south-west Ethiopia’s dwindling tropical rain-forest." The Majangir language is part of the Surmic cluster, however it is the most isolated language in that cluster ( Harold C. Fleming 1983). A language survey has shown that dialect variation from north to south

5250-592: The Northern Hemisphere, colonizing Europe from Africa by the Paleocene , and then spreading east to Asia . This was facilitated by the warming climate around the same time, allowing bees to move to higher latitudes following the spread of tropical and subtropical habitats. By the Eocene (~45 mya) there was already considerable diversity among eusocial bee lineages. A second extinction event among bees

5355-580: The United States, according to the National Honey Board, "Ensuring honey authenticity is one of the great challenges facing the honey industry today. Over the past half century, a number of honey testing methods have been developed to detect food fraud. To date, there is no single universal analytical method available which is capable of detecting all types of adulteration with adequate sensitivity." Isotope ratio mass spectrometry can be used to detect addition of corn syrup and cane sugar by

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5460-430: The ability to absorb moisture directly from the air, a phenomenon called hygroscopy . The amount of water the honey absorbs is dependent on the relative humidity of the air. Because honey contains yeast, this hygroscopic nature requires that honey be stored in sealed containers to prevent fermentation, which usually begins if the honey's water content rises much above 25%. Honey tends to absorb more water in this manner than

5565-555: The adulteration from buyers until the honey was liquefied. In modern times, the most common adulterant became clear, almost-flavorless corn syrup; the adulterated mixture can be very difficult to distinguish from pure honey. According to the Codex Alimentarius of the United Nations, any product labeled as "honey" or "pure honey" must be a wholly natural product, although labeling laws differ between countries. In

5670-488: The appearance of being social. Large groups of solitary bee nests are called aggregations , to distinguish them from colonies . In some species, multiple females share a common nest, but each makes and provisions her own cells independently. This type of group is called "communal" and is not uncommon. The primary advantage appears to be that a nest entrance is easier to defend from predators and parasites when multiple females use that same entrance regularly. The life cycle of

5775-438: The available honey and replace the entire colony the next spring. Since the invention of removable frames, the principles of husbandry led most beekeepers to ensure that their bees have enough stores to survive the winter, either by leaving some honey in the beehive or by providing the colony with a honey substitute such as sugar water or crystalline sugar (often in the form of a "candyboard"). The amount of food necessary to survive

5880-848: The bee families is based on Hedtke et al., 2013, which places the former families Dasypodaidae and Meganomiidae as subfamilies inside the Melittidae. English names, where available, are given in parentheses. Melittidae (inc. Dasypodainae , Meganomiinae ) at least 50 Mya [REDACTED] Apidae (inc. honeybees, cuckoo bees, carpenter bees) ≈87 Mya [REDACTED] Megachilidae (mason, leafcutter bees) ≈50 Mya [REDACTED] Andrenidae (mining bees) ≈34 Mya [REDACTED] Halictidae (sweat bees) ≈50 Mya [REDACTED] Colletidae (plasterer bees) ≈25 Mya [REDACTED] Stenotritidae (large Australian bees) ≈2 Mya [REDACTED] Bees differ from closely related groups such as wasps by having branched or plume-like setae (hairs), combs on

5985-677: The best known insects. Their colonies are established by swarms , consisting of a queen and several thousand workers. There are 29 subspecies of one of these species, Apis mellifera , native to Europe, the Middle East, and Africa. Africanized bees are a hybrid strain of A. mellifera that escaped from experiments involving crossing European and African subspecies; they are extremely defensive. Stingless bees are also highly eusocial . They practise mass provisioning , with complex nest architecture and perennial colonies also established via swarming. Many bumblebees are eusocial, similar to

6090-481: The carbon isotopic signature . Addition of sugars originating from corn or sugar cane ( C4 plants , unlike the plants used by bees, and also sugar beet , which are predominantly C3 plants ) skews the isotopic ratio of sugars present in honey, but does not influence the isotopic ratio of proteins. In an unadulterated honey, the carbon isotopic ratios of sugars and proteins should match. Levels as low as 7% of addition can be detected. In 2020, global production of honey

6195-674: The case of the best-known bee species, the western honey bee , for producing honey . Bees are a monophyletic lineage within the superfamily Apoidea . They are currently considered a clade , called Anthophila . There are over 20,000 known species of bees in seven recognized biological families . Some species – including honey bees , bumblebees , and stingless bees  – live socially in colonies while most species (>90%) – including mason bees , carpenter bees , leafcutter bees , and sweat bees  – are solitary. Bees are found on every continent except Antarctica , in every habitat on

6300-427: The colony to a site for a new hive. Bees generally swarm before a suitable location for another hive has been discovered by scouts sent out for this purpose. Until such a location is found the swarm will simply conglomerate near the former hive, often from tree branches. These swarms are unusually docile and amenable to transport by humans. When provided with a suitable nesting site, such as a commercial Langstroth hive ,

6405-401: The decline in wild bees has increased the value of pollination by commercially managed hives of honey bees. The analysis of 353 wild bee and hoverfly species across Britain from 1980 to 2013 found the insects have been lost from a quarter of the places they inhabited in 1980. Human beekeeping or apiculture ( meliponiculture for stingless bees) has been practised for millennia, since at least

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6510-530: The end of the Ethiopian Civil War in 1991, with its subsequent remapping of Ethiopia by ethnic lines, the Majangir have felt very marginalized politically, their territory now divided among three kalil or administrative Regions (Sato 2000 and 2002). This frustration has led to some armed fighting with the government (Vaughan 2003:268). Currently, many outsiders are buying or leasing land from

6615-487: The eusocial Vespidae such as hornets in that the queen initiates a nest on her own rather than by swarming. Bumblebee colonies typically have from 50 to 200 bees at peak population, which occurs in mid to late summer. Nest architecture is simple, limited by the size of the pre-existing nest cavity, and colonies rarely last more than a year. In 2011, the International Union for Conservation of Nature set up

6720-480: The family Crabronidae , which were predators of other insects. The switch from insect prey to pollen may have resulted from the consumption of prey insects which were flower visitors and were partially covered with pollen when they were fed to the wasp larvae. This same evolutionary scenario may have occurred within the vespoid wasps, where the pollen wasps evolved from predatory ancestors. Based on phylogenetic analysis, bees are thought to have originated during

6825-530: The females emerge. Solitary bees are very unlikely to sting (only in self-defense, if ever), and some (esp. in the family Andrenidae ) are stingless. While solitary, females each make individual nests. Some species, such as the European mason bee Hoplitis anthocopoides , and the Dawson's Burrowing bee , Amegilla dawsoni, are gregarious, preferring to make nests near others of the same species, and giving

6930-605: The first appearance of bees. The novelty is that bees are specialized as pollination agents, with behavioral and physical modifications that specifically enhance pollination, and are the most efficient pollinating insects. In a process of coevolution , flowers developed floral rewards such as nectar and longer tubes, and bees developed longer tongues to extract the nectar. Bees also developed structures known as scopal hairs and pollen baskets to collect and carry pollen. The location and type differ among and between groups of bees. Most species have scopal hairs on their hind legs or on

7035-497: The flora also have an effect on honey aroma properties, such as a "fruity" or "grassy" aroma from longan honey, or a "waxy" aroma from sunflower honey. Dominant VOCs in one study were linalool oxide , trans-linalool oxide, 2- phenylacetaldehyde , benzyl ethanol , isophorone , and methyl nonanoate . VOCs can also be introduced from the bodies of the bees, be produced by the enzymatic actions of digestion, or from chemical reactions that occur between different substances within

7140-470: The forelimbs for cleaning their antennae, small anatomical differences in limb structure, and the venation of the hind wings; and in females, by having the seventh dorsal abdominal plate divided into two half-plates. Bees have the following characteristics: The largest species of bee is thought to be Wallace's giant bee Megachile pluto , whose females can attain a length of 39 millimetres (1.54 in). The smallest species may be dwarf stingless bees in

7245-460: The fossil record, all modern bee families had already diverged from one another by the end of the Cretaceous. The Melittidae , Apidae , and Megachilidae had already evolved on the supercontinent prior to its fragmentation. Further divergences were facilitated by West Gondwana's breakup around 100 million years ago, leading to a deep Africa-South America split within both the Apidae and Megachilidae,

7350-464: The group consists of a mother (the queen ) and her daughters ( workers ). When the castes are purely behavioural alternatives, with no morphological differentiation other than size, the system is considered primitively eusocial, as in many paper wasps ; when the castes are morphologically discrete, the system is considered highly eusocial. True honey bees (genus Apis , of which eight species are currently recognized) are highly eusocial, and are among

7455-475: The hive and after its removal by a beekeeper . Honey bees are not the only eusocial insects to produce honey. All non-parasitic bumblebees and stingless bees produce honey. Some wasp species, such as Brachygastra lecheguana and Brachygastra mellifica , found in South and Central America, are known to feed on nectar and produce honey. Other wasps, such as Polistes versicolor , also consume honey. In

7560-425: The hive, through water evaporation that concentrates the honey's sugars until it is thick and viscous. Honey bees stockpile honey in the hive . Within the hive is a structure made from wax called honeycomb . The honeycomb is made up of hundreds or thousands of hexagonal cells, into which the bees regurgitate honey for storage. Other honey-producing species of bee store the substance in different structures, such as

7665-466: The honey during storage, and therefore may change, increase, or decrease over long periods of time. VOCs may be produced, altered, or greatly affected by temperature and processing. Some VOCs are heat labile, and are destroyed at elevated temperatures, while others can be created during non-enzymatic reactions, such as the Maillard reaction . VOCs are responsible for nearly all of the aroma produced by

7770-404: The honey stomach holds about 40 mg of liquid. This is about half the weight of an unladen bee. Collecting this quantity in nectar can require visits to more than a thousand flowers. When nectar is plentiful, it can take a bee more than an hour of ceaseless work to collect enough nectar to fill its honey crop. Salivary enzymes and proteins from the bee's hypopharyngeal gland are secreted into

7875-431: The honey-storage areas. This temperature is regulated either by generating heat with their bodies or removing it through water evaporation. The evaporation removes water from the stored honey, drawing heat from the colony. The bees use their wings to govern hive cooling. Coordinated wing beating moves air across the wet honey, drawing out water and heat. Ventilation of the hive eventually expels both excess water and heat into

7980-447: The individual sugars allow on their own, which may be due to other ingredients it contains. Fermentation of honey usually occurs after crystallization, because without the glucose, the liquid portion of the honey primarily consists of a concentrated mixture of fructose, acids, and water, providing the yeast with enough of an increase in the water percentage for growth. Honey that is to be stored at room temperature for long periods of time

8085-607: The isolation of the Melittidae in Africa, and the origins of the Colletidae , Andrenidae and Halictidae in South America. The rapid radiation of the South American bee families is thought to have followed the concurrent radiation of flowering plants in the same region. Later in the Cretaceous (80 million years ago), colletid bees colonized Australia from South America (with an offshoot lineage evolving into

8190-661: The lineage as incertae sedis within the Aculeata . The Allodapini (within the Apidae) appeared around 53 Mya. The Colletidae appear as fossils only from the late Oligocene (~25 Mya) to early Miocene . The Melittidae are known from Palaeomacropis eocenicus in the Early Eocene . The Megachilidae are known from trace fossils (characteristic leaf cuttings) from the Middle Eocene . The Andrenidae are known from

8295-488: The male line. A person cannot marry a person from the same clan ( exogamy ), nor should they marry a person from their mother's clan (Stauder 1971, Unseth 1998a). The Majangir traditionally made two kinds of alcoholic drink: one from grain "tááján" (cf. tella ) and one from honey "ògòòl" (cf. tej ; Teramoto et al., 2005). The Majangir have traditionally used a number of musical instruments, sometimes to accompany singing and sometimes played without. Their instruments include

8400-494: The middle of their life cycles they alternate between feeding on protein-rich pollen and feeding on honey, which is a far denser source of food energy . Human beings have semi-domesticated several species of honey bee by taking advantage of their swarming stage. Swarming is the means by which new colonies are established when there is no longer space for expansion in the colony's present hive. The old queen lays eggs that will develop into new queens and then leads as many as half

8505-415: The minor substances in honey can be affected greatly by heating, changing the flavor, aroma, or other properties, so heating is usually done at the lowest temperature and for the shortest time possible. The average pH of honey is 3.9, but can range from 3.4 to 6.1. Honey contains many kinds of acids, both organic and amino . However, the different types and their amounts vary considerably, depending on

8610-567: The mixture of VOCs in different honeys in one review, longan honey had a higher amount of volatiles (48 VOCs), while sunflower honey had the lowest number of volatiles (8 VOCs). VOCs are primarily introduced into the honey from the nectar, where they are excreted by the flowers imparting individual scents. The specific types and concentrations of certain VOCs can be used to determine the type of flora used to produce monofloral honeys. The specific geography, soil composition and acidity used to grow

8715-501: The multiple (at least nine) evolutions of eusociality within Hymenoptera . Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such as termites are not haplodiploid. Conversely, all bees are haplodiploid but not all are eusocial, and among eusocial species many queens mate with multiple males, creating half-sisters that share only 25% of each other's genes. But, monogamy (queens mating singly)

8820-565: The necessary adaptations to do this. However, certain wasp species such as pollen wasps have similar behaviours, and a few species of bee scavenge from carcases to feed their offspring. Solitary bees are important pollinators; they gather pollen to provision their nests with food for their brood. Often it is mixed with nectar to form a paste-like consistency. Some solitary bees have advanced types of pollen-carrying structures on their bodies. Very few species of solitary bee are being cultured for commercial pollination. Most of these species belong to

8925-402: The nectar once it is in the bee's honey stomach. These substances begin cleaving complex sugars like sucrose and starches into simpler sugars such as glucose and fructose . This process slightly raises the water content and the acidity of the partially digested nectar. Once filled, the forager bees return to the hive. There they regurgitate and transfer nectar to hive bees. Once it

9030-430: The other members of Hymenoptera bees are haplodiploid ; the sex of a bee is determined by whether or not the egg is fertilised. After mating, a female stores the sperm, and determines which sex is required at the time each individual egg is laid, fertilised eggs producing female offspring and unfertilised eggs, males. Tropical bees may have several generations in a year and no diapause stage. Honey Honey

9135-402: The outside world. The process of evaporating continues until the honey reaches its final water content of between 15.5% to 18%. This concentrates the sugars far beyond the saturation point of water, which is to say there is far more sugar dissolved in what little water remains in honey than ever could be dissolved in an equivalent volume of water. Honey, even at hive temperatures, is therefore

9240-547: The planet that contains insect-pollinated flowering plants . The most common bees in the Northern Hemisphere are the Halictidae , or sweat bees, but they are small and often mistaken for wasps or flies. Bees range in size from tiny stingless bee species, whose workers are less than 2 millimetres (0.08 in) long, to the leafcutter bee Megachile pluto , the largest species of bee, whose females can attain

9345-456: The polarization plane. The fructose gives a negative rotation, while the glucose gives a positive one. The overall rotation can be used to measure the ratio of the mixture. Honey is generally pale yellow and dark brown in color, but other colors can occur, depending on the sugar source. Bee colonies that forage on Kudzu ( Pueraria montana var. lobata ) flowers, for example, produce honey that varies in color from red to purple. Honey has

9450-456: The pots made of wax and resin used by the stingless bee . Honey for human consumption is collected from wild bee colonies, or from the hives of domesticated bees. The honey produced by honey bees is the most familiar to humans, thanks to its worldwide commercial production and availability. The husbandry of bees is known as beekeeping or apiculture, with the cultivation of stingless bees usually referred to as meliponiculture . Honey

9555-431: The processed nectar to bubbling and then passing the refined liquid on to others. It can take as long as 20 minutes of continuous regurgitation, digestion and evaporation until the product reaches storage quality. The new honey is then placed in honeycomb cells, which are left uncapped. This honey still has a very high water content, up to 70%, depending on the concentration of nectar gathered. At this stage of its refinement

9660-415: The quality of honey in terms of ash content. The effect honey has on light is useful for determining the type and quality. Variations in its water content alter its refractive index . Water content can easily be measured with a refractometer . Typically, the refractive index for honey ranges from 1.504 at 13% water content to 1.474 at 25%. Honey also has an effect on polarized light , in that it rotates

9765-435: The reaction can also be slowed by storing the honey at colder temperatures. Unlike many other liquids, honey has very poor thermal conductivity of 0.5 W/(m⋅K) at 13% water content (compared to 401 W/(m⋅K) of copper ), taking a long time to reach thermal equilibrium . Due to its high kinematic viscosity honey does not transfer heat through momentum diffusion ( convection ) but rather through thermal diffusion (more like

9870-509: The role of "tapad" (final implosive d), who served as ritual leaders. These were from the Meelanir clan, a group has links with similar-named privileged clans in other Surmic groups (Unseth and Abbink 1998). In case of a serious disagreement, one party would simply move away. There was no standard social reconciliation mechanism as is found in highland Ethiopian cultures. The Majangir have over 70 clans, with clan identity passed down through

9975-503: The sap on which those insects feed and is usually somewhat more dilute than nectar. One source describes the water content of honeydew as around 89%. Whether it is feeding on nectar or honeydew, the bee sucks these runny fluids through its proboscis , which delivers the liquid to the bee's honey stomach or "honey crop". This cavity lies just above its food stomach, the latter of which digests pollen and sugars consumed by an individual honey bee for its own nourishment. In Apis mellifera ,

10080-441: The swarm will readily form a new colony in artificial surroundings. These semi-domesticated colonies are then looked after by humans practicing apiculture or meliponiculture. Captured bees are encouraged to forage, often in agricultural settings such as orchards, where pollinators are highly valued. The honey, pollen , wax and resins the bees produce are all harvested by humans for a variety of uses. The term "semi-domesticated"

10185-492: The times of Ancient Egypt and Ancient Greece . Bees have appeared in mythology and folklore, through all phases of art and literature from ancient times to the present day, although primarily focused in the Northern Hemisphere where beekeeping is far more common. In Mesoamerica , the Mayans have practiced large-scale intensive meliponiculture since pre-Columbian times. The immediate ancestors of bees were stinging wasps in

10290-604: The tribe Meliponini whose workers are less than 2 millimetres (0.08 in) in length. According to inclusive fitness theory, organisms can gain fitness not just through increasing their own reproductive output, but also that of close relatives. In evolutionary terms, individuals should help relatives when Cost < Relatedness * Benefit . The requirements for eusociality are more easily fulfilled by haplodiploid species such as bees because of their unusual relatedness structure. In haplodiploid species, females develop from fertilized eggs and males from unfertilized eggs. Because

10395-425: The type of flora used to produce it (pasturage), temperature, and the proportion of the specific sugars it contains. Fresh honey is a supersaturated liquid, containing more sugar than the water can typically dissolve at ambient temperatures. At room temperature, honey is a supercooled liquid, in which the glucose precipitates into solid granules. This forms a semisolid solution of precipitated glucose crystals in

10500-717: The type of honey. These acids may be aromatic or aliphatic (nonaromatic). The aliphatic acids contribute greatly to the flavor of honey by interacting with the flavors of other ingredients. Organic acids comprise most of the acids in honey, accounting for 0.17–1.17% of the mixture, with gluconic acid formed by the actions of glucose oxidase as the most prevalent. Minor amounts of other organic acids are present, consisting of formic , acetic , butyric , citric , lactic , malic , pyroglutamic , propionic , valeric , capronic , palmitic , and succinic , among many others. Individual honeys from different plant sources contain over 100 volatile organic compounds (VOCs), which play

10605-501: The underside of their abdomens. Some species in the family Apidae have pollen baskets on their hind legs, while very few lack these and instead collect pollen in their crops. The appearance of these structures drove the adaptive radiation of the angiosperms , and, in turn, bees themselves. Bees coevolved not only with flowers but it is believed that some species coevolved with mites. Some provide tufts of hairs called acarinaria that appear to provide lodgings for mites; in return, it

10710-500: The water content of the honey is high enough that ubiquitous yeast spores can reproduce in it, a process which, if left unchecked, would rapidly consume the new honey's sugars. To combat this, bees use an ability rare among insects: the endogenous generation of heat. Bees are among the few insects that can create large amounts of body heat. They use this ability to produce a constant ambient temperature in their hives. Hive temperatures are usually around 35 °C (95 °F) in

10815-613: The winter depends on the variety of bees and on the length and severity of local winters. Many animal species are attracted to wild or domestic sources of honey. Because of its composition and chemical properties, honey is suitable for long-term storage, and is easily assimilated even after long preservation. Honey, and objects immersed in honey, have been preserved for centuries. (However, no edible honey has been found in Egyptian tombs; all such cases have been proven to be other substances or only chemical traces. ) The key to preservation

10920-401: Was 1.8   million tonnes , led by China with 26% of the world total (table). Other major producers were Turkey , Iran , Argentina , and Ukraine . Over its history as a food, the main uses of honey are in cooking, baking, desserts, as a spread on bread, as an addition to various beverages such as tea, and as a sweetener in some commercial beverages. Due to its energy density, honey

11025-895: Was not included in this analysis. Further studies by Sann et al. , 2018, elevated the subfamilies (plus one tribe and one subtribe) of Crabronidae sensu lato to family status. They also recovered the placement of Heterogyna within Nyssonini and sunk Heterogynaidae . The newly erected family, Ammoplanidae , formerly a subtribe of Pemphredoninae , was recovered as the most sister family to bees. Ampulicidae (Cockroach wasps) [REDACTED] Astatidae [REDACTED] Bembicidae [REDACTED] Sphecidae ( sensu stricto ) [REDACTED] Crabronidae ( sensu stricto ) [REDACTED] Mellinidae [REDACTED] Pemphredonidae [REDACTED] Philanthidae [REDACTED] Psenidae [REDACTED] Ammoplanidae Anthophila (bees) [REDACTED] This cladogram of

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