Bee

Bees are flying closely related to and , known for their role in and, in the case of the best-known bee species, the , for producing and . Bees are a lineage within the superfamily . They are presently considered a , called Anthophila. There are over 16,000 known species of bees in seven recognized . They are found on every continent except for , in every habitat on the planet that contains insect-pollinated .

Some species — including , , and — live in colonies. Bees are adapted for feeding on and , the former primarily as an energy source and the latter primarily for and other nutrients. Most pollen is used as food for . Bee is important both ecologically and . The decline in wild bees has increased the value of pollination by commercially managed hives of honey bees.

Bees range in size from tiny species whose workers are less than 2 millimetres (0.08 in) long, to , the largest species of , whose females can attain a length of 39 millimetres (1.54 in). The most common bees in the are the , or sweat bees, but they are small and often mistaken for wasps or flies. Vertebrate predators of bees include birds such as ; insect predators include and .

Human or apiculture has been practised for millennia, since at least the times of and . Apart from honey and pollination, honey bees produce beeswax, and . Bees have appeared in mythology and folklore, through all phases of art and literature, from ancient times to the present day, though primarily focused in the , where is far more common.

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.[3]

The ancestors of bees were in the family , which were 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 scenario may have occurred within the wasps, where the evolved from predatory ancestors. Until recently, the oldest non-compression bee fossil had been found in New Jersey , Cretotrigona prisca of , a . (~100 mya), , is considered "an extinct lineage of pollen-collecting Apoidea to the modern bees".) place it clearly within the bees, but it retains two unmodified ancestral traits () of the legs (two mid-tibial spurs, and a slender hind basitarsus), showing its transitional status. (~45 mya) there was already considerable diversity among eusocial bee lineages.

The highly appeared roughly 87 Mya, and the (within the Apidae) around 53 Mya.[9]
The appear as fossils only from the late (~25 Mya) to early .[10]
The are known from Palaeomacropis eocenicus in the .[11]
The are known from trace fossils (characteristic leaf cuttings) from the .[12]
The are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale.[13]
The first appear in the Early Eocene are known from fossil brood cells of age.[17]

Coevolution

The earliest animal-pollinated flowers were shallow, cup-shaped blooms by insects such as , so the was well established before the first appearance of bees. The novelty is that bees are as pollination agents, with behavioral and physical modifications that specifically enhance pollination, and are the most efficient pollinating insects. In a process of , flowers developed floral rewards and longer tubes, and bees developed longer tongues to extract the nectar. and 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 the underside of their abdomens. Some species in the family Apidae have on their hind legs, while very few lack these and instead collect pollen in their crops. of the , and, in turn, bees themselves. 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 is believed that mites eat fungi that attack pollen, so the relationship in this case may be .

Phylogeny

External

This is based on Debevic et al, 2012, which used molecular phylogeny to demonstrate that the bees () arose from deep within the , which is therefore . The placement of the is uncertain. was not included in this analysis.

Internal

This cladogram of the bee families is based on Hedtke et al., 2013, which places the former families Dasypodaidae and Meganomiidae as subfamilies inside the Melittidae.[23] English names, where available, are given in parentheses.

Bees differ from closely related groups such as wasps by having branched or plume-like (hairs), combs on 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.[24]

Bees have the following characteristics:

• A pair of large which cover much of the surface of the head. Between and above these are three small simple eyes () which provide information on light intensity.


• The usually have 13 segments in males and 12 in females, and are , having an elbow joint part way along. They house large numbers of sense organs that can detect touch (mechanoreceptors), smell and taste; and small, hairlike mechanoreceptors that can detect air movement so as to "hear" sounds.


• The are adapted for both chewing and sucking by having both a pair of and a long for sucking up nectar.[25]


• The thorax has three segments, each with a pair of robust legs, and a pair of membranous wings on the hind two segments. The front legs of corbiculate bees bear combs for cleaning the antennae, and in many species the hind legs bear pollen baskets, flattened sections with incurving hairs to secure the collected pollen. The wings are synchronised in flight, and the somewhat smaller hind wings connect to the forewings by a row of hooks along their margin which connect to a groove in the forewing.


• The abdomen has nine segments, the hindermost three being modified into the sting., whose females can attain a length of 39 millimetres (1.54 in). whose workers are less than 2 millimetres (0.08 in) in length.[27]
  
  
  Sociality
  

  Haplodiploid breeding system

  According to 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 species such as bees because of their unusual relatedness structure. (has only one copy of each gene), his daughters (which are , 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 termed "supersisters", more closely related to their sisters than they would be to their own offspring..

  Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such as 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.
  

  
  

  Eusociality

  
  

  Bees may be solitary or may live in various types of communities. appears to have originated from at least three independent origins in halictid bees. colonies; these are characterised by cooperative brood care and a into reproductive and non-reproductive adults, plus overlapping generations.. In some species, groups of cohabiting females may be sisters, and if there is a division of labour within the group, they are considered . The group is called eusocial if, in addition, the group consists of a mother (the ) and her daughters (). When the castes are purely behavioural alternatives, with no morphological differentiation other than size, the system is considered primitively eusocial, as in many ; when the castes are morphologically discrete, the system is considered highly eusocial.[19]
  

  True honey bees (genus , of which seven species are currently recognized) are highly eusocial, and are among the best known insects. Their colonies are established by , consisting of a queen and several hundred workers. There are 29 subspecies of one of these species, , native to Europe, the Middle East, and Africa. are a hybrid strain of A. mellifera that escaped from experiments involving crossing European and African subspecies; they are extremely defensive.[35]
  

  are also highly . They practise , with complex nest architecture and perennial colonies also established via swarming.[36]
  

  
  A carrying pollen in its (corbiculae)
  

  Many bumblebees are eusocial, similar to the eusocial such as 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. set up the Bumblebee Specialist Group to review the threat status of all bumblebee species worldwide using the criteria.[38]
  

  There are many more species of primitively eusocial than highly eusocial bees, but they have been studied less often. Most are in the family , 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 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 . or have a mix of eusocial and solitary nests in the same population. (Apidae) include some primitively eusocial species with similar biology. Some bees (Apidae) form primitively eusocial colonies, with : a larva's food is supplied gradually as it develops, as is the case in honey bees and some bumblebees.[42]
  

  
  

  Solitary and communal bees

  
  A leafcutting bee, , cutting circles from acacia leaves
  

  Most other bees, including familiar insects such as , and 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 . Bees collect pollen to feed their young, and have the necessary adaptations to do this. However, certain wasp species such as have similar behaviours, and a few species of bee from carcases to feed their offspring. which are commonly known by their nesting behavior or preferences, namely: carpenter bees, , mason bees, , , , leafcutter bees, and .[43]
  

  Most solitary bees nest in the ground in a variety of soil textures and conditions while others create nests in hollow or twigs, holes in . The female typically creates 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 the females emerge. Solitary bees are either stingless or very unlikely to sting (only in self-defense, if ever).
  

  While solitary, females each make individual nests. Some species, such as the European mason bee ,, Amegilla dawsoni,. 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 there are multiple females using that same entrance on a regular basis.[46]
  

  
  Biology
  

  Life cycle

  The life cycle of a bee, be it a solitary or social species, involves the laying of an egg, the development through several moults of a legless , a stage during which the insect undergoes , followed by the emergence of a winged adult. 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. 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 stage.
  

  The egg is generally oblong, slightly curved and tapering at one end. Solitary bees, lay each egg in a separate cell with a supply of mixed pollen and nectar next to it. This may be rolled into a pellet or placed in a pile and is known as mass provisioning. Social bee species provision progressively, that is, they feed the larva regularly while it grows. The nest varies from a hole in the ground or in wood, in solitary bees, to a substantial structure with wax combs in bumblebees and honey bees.[52]
  

  In most species, larvae are whitish grubs, roughly oval and bluntly-pointed at both ends. They have 15 segments and in each segment for breathing. They have no legs but move within the cell, helped by tubercles on their sides. They have short horns on the head, jaws for chewing food and an appendage on either side of the mouth tipped with a bristle. There is a gland under the mouth that secretes a viscous liquid which solidifies into the silk they use to produce a cocoon. The cocoon is semi-transparent and the pupa can be seen through it. Over the course of a few days, the larva undergoes metamorphosis into an winged adult. When ready to emerge, the adult splits its skin dorsally and climbs out of the and breaks out of the cell.[52]
  

  Flight

  
  Honeybee in flight carrying pollen in

  's 1934 book Le vol des insectes, says that he and had applied the equations of to and found that their flight could not be explained by fixed-wing calculations, but that "One shouldn't be surprised that the results of the calculations don't square with reality".. High-speed showed that lift was generated by "the unconventional combination of short, choppy wing strokes, a rapid rotation of the wing as it flops over and reverses direction, and a very fast wing-beat frequency". Wing-beat frequency normally increases as size decreases, but as the bee's wing beat covers such a small , it flaps approximately 230 times per second, faster than a (200 times per second) which is 80 times smaller.[58]
  

  
  

  Navigation, communication, and finding food

  The ethologist studied in the honey bee. He showed that honey bees communicate by the , in which a worker indicates the location of a food source to other workers in the hive. He demonstrated that bees can recognize a desired compass direction in three different ways: by the sun, by the pattern of the blue sky, and by the earth's magnetic field. He showed that the sun is the preferred or main compass; the other mechanisms are used under cloudy skies or inside a dark .
  

  
  

  Digestion

  
  

  The gut of bees is relatively simple, but multiple metabolic strategies exist in the gut . sugars and so easily absorbed, pollen contains complex : branching and . and two groups of ), and two other groups in complex sugars ( and ). Digestion of pectin and hemicellulose is dominated by bacterial Gilliamella and respectively. Bacteria that cannot digest polysaccharides obtain enzymes from their neighbors, and bacteria that lack certain amino acids do the same, creating multiple .[63]
  

  Although most bee species are and , some are not. Particularly unusual are in the genus , which consume carrion and wasp brood, turning meat into a honey-like substance.[64]
  

  
  Ecology
  

  Floral relationships

  
  

  Most bees are polylectic (generalist) meaning they collect pollen from a range of flowering plants, but some are (specialists), in that they only gather pollen from one or a few species or genera of closely related plants. (one of the few cases where male bees are effective pollinators). Bees are able to sense the presence of desirable flowers through ultraviolet patterning on flowers, floral odors, Once landed, a bee then uses nectar quality to determine whether to continue visiting similar flowers.
  

  In rare cases, a species may only be effectively pollinated by a single bee species, and some plants are at least in part because their pollinator is also threatened. But, there is a pronounced tendency for oligolectic bees to be associated with common, widespread plants visited by multiple pollinator species. For example, the in the arid parts of the United States southwest is associated with some 40 oligoleges.[69]
  

  
  

  As mimics and models

  lures male bees to attempt to mate with the flower's lip, which resembles a bee perched on a pink flower.
  

  Many bees are coloured, typically orange and black, warning of their ability to defend themselves with a powerful sting. As such they are models for by non-stinging insects such as , and ,
  

  Bees are themselves of other aposematic insects with the same colour scheme, including , and other beetles, and many butterflies and moths () which are themselves distasteful, often through acquiring bitter and poisonous chemicals from their plant food. All the Müllerian mimics, including bees, benefit from the reduced risk of predation that results from their easily recognised warning coloration.[71]
  

  Bees are also mimicked by plants such as the which imitates both the appearance and the scent of a female bee; male bees attempt to mate () with the furry lip of the flower, thus pollinating it.[72]
  

  
  

  As brood parasites

  
  , a of the bumblebee

  occur in several bee families including the subfamily .) and do not construct their own nests. They typically enter the nests of pollen collecting species, and lay their eggs in cells provisioned by the host bee. When the "cuckoo" bee larva hatches, it consumes the host larva's pollen ball, and often the host egg also. is an aggressive species that attacks and enslaves other bees of the same subgenus. However, unlike many other bee brood parasites, they have pollen baskets and often collect pollen.[75]
  

  In Southern Africa, hives of African honeybees (A. mellifera scutellata) are being destroyed by parasitic workers of the Cape honeybee, A. m. capensis. These lay eggs (""), escaping normal , leading to the colony's destruction; the parasites can then move to other hives.[76]
  

  The in the subgenus Psithyrus are closely related to, and resemble, their hosts in looks and size. This common pattern gave rise to the ecological principle "". Others parasitize bees in different families, like , a , two species of which are cleptoparasites of the genus , bees.[78]
  

  
  

  Nocturnal bees

  
  

  Four bee families (, , , and ) contain some species that are . Most are tropical or subtropical, but some live in arid regions at higher latitudes. These bees have greatly enlarged , which are extremely sensitive to light and dark, though incapable of forming images. Some have refracting superposition compound eyes: these combine the output of many elements of their compound eyes to provide enough light for each retinal photoreceptor. Their ability to fly by night enables them to avoid many predators, and to exploit flowers that produce nectar only or also at night.[79]
  

  
  

  Predators, parasites and pathogens

  The bee-eater, , specialises in feeding on bees; here a male catches a nuptial gift for his mate.
  

  Vertebrate predators of bees include , and , which make short sallies to catch insects in flight. fly almost continually, catching insects as they go. The attacks bees' nests and eats the larvae. interacts with humans by guiding them to the nests of wild bees. The humans break open the nests and take the honey and the bird feeds on the larvae and the wax. dig up bumblebee nests and eat both the larvae and any stored food.[83]
  

  
  The beewolf paralysing a bee with its sting
  

  Specialist ambush predators of visitors to flowers include , which wait on flowering plants for pollinating insects; , and , of the tropics) wait motionless, camouflaged as flowers. are large wasps that habitually attack bees; estimated that a single colony of the beewolf might kill several thousand honeybees in a day: all the prey he observed were honeybees. and . and .
  

  
  Relationship with humans
  

  In mythology and folklore

  Gold plaques embossed with winged bee goddesses. , . 7th century B.C.
  

  's describes three bee-maidens with the power of and thus speaking truth, and identifies the food of the gods as honey. Sources associated the bee maidens with and, until the 1980s, scholars followed Gottfried Hermann (1806) in incorrectly identifying the bee-maidens with the .. Bees were also associated with the and the prophetess was sometimes called a bee.[88]
  

  The image of a community of honey bees has been used from ancient to modern times, in and ; in and ; in and ; , and by political and social theorists such as and as a model for human .".[90]
  

  
  

  In art and literature

  Some of the oldest examples of bees in art are in which have been dated to 15,000 BC.[91]
  

  's poem (1888) contains the couplet "Nine bean rows will I have there, a hive for the honey bee, / And live alone in the bee loud glade." At the time he was living in in the West of London.'s illustrated book (1910) features Babbity Bumble and her brood (pictured). ' book (1984) uses bees and beekeeping as part of its story and puzzle. 's (2004), and the starring , tells the story of a girl who escapes her abusive home and finds her way to live with a family of beekeepers, the Boatwrights.
  

  The humorous 2007 animated film used 's first script and was his first work for children; he starred as a bee named Barry B. Benson, alongside . Critics found its premise awkward and its delivery tame.'s (2014) describes his efforts to save bumblebees in Britain, as well as much about their biology. The playwright Laline Paull's The Bees (2015) tells the tale of a hive bee named Flora 717 from hatching onwards.[94]
  

  
  

  Beekeeping

  A commercial beekeeper at work
  

  Humans have kept honey bee colonies, commonly in , for millennia. collect , , , , and from hives; bees are also kept to crops and to produce bees for sale to other beekeepers.
  

  Depictions of humans collecting honey from wild bees date to 15,000 years ago; efforts to domesticate them are shown in Egyptian art around 4,500 years ago. such as . From the 18th century, European understanding of the colonies and biology of bees allowed the construction of the moveable comb hive so that honey could be harvested without destroying the colony. Among Classical Era authors, beekeeping with the use of smoke is described in Aristotle's Book 9., but "kings" rather than queens; predators including toads and bee-eaters; and the , with the "irresistible suggestion" of άpοσειονται ("aroseiontai", it waggles) and παρακολουθούσιν ("parakolouthousin", they watch).
  

  Beekeeping is described in detail by in his ; it is also mentioned in his , and in Pliny's .[101]
  

  
  

  As commercial pollinators

  (Apidae) are important pollinators of and .
  
  

  Bees play an important role in , and are the major type of in many that contain flowering plants. It is estimated that one third of the human food supply depends on by insects, birds and bats, most of which is accomplished by bees, whether wild or domesticated. Over the last half century, there has been a general decline in the species richness of wild bees and other pollinators, probably attributable to stress from increased parasites and disease, the use of pesticides, and a general decrease in the number of wild flowers. Climate change probably exacerbates the problem.[104]
  

  has overtaken the role of honey production for in many countries. After the introduction of , honey bees declined dramatically in the US, though their numbers have since recovered. The number of colonies kept by beekeepers declined slightly, through , systematic pesticide use, and mites, and the closure of beekeeping businesses. In 2006 and 2007 the rate of attrition increased, and was described as . were shown to be in every killed colony, and deadly in combination. Winter losses increased to about 1/3. Varroa mites were thought to be responsible for about half the losses.[114]
  

  Apart from colony collapse disorder, losses outside the US have been attributed to causes including pesticide seed dressings, using such as , and . From 2013 the restricted some pesticides to stop bee populations from declining further. report warned that bees faced increased risk of extinction because of .
  

  Farmers have focused on alternative solutions to mitigate these problems. By raising native plants, they provide food for native bee pollinators like ,[121] leading to less reliance on honey bee populations.
  

  
  

  As food producers

  
  

  Honey is a natural product produced by bees and stored for their own use, but its sweetness has always appealed to humans. Before domestication of bees was even attempted, humans were raiding their nests for their honey. Smoke was often used to subdue the bees and such activities are depicted in in Spain dated to 15,000 BC.[91]
  

  Honey bees are used commercially to produce . ,,.
  

  
  

  As food (bee brood)

  
  Bee larvae as food in the dish
  

  Bees are partly considered . Indigenous people in many countries , including the larvae and pupae of bees, mostly stingless species. They also gather larvae, pupae and surrounding cells, known as , for consumption. dish from Central and East , bee larvae are eaten as a companion to , after being mixed with shredded , wrapped in leaves, and steamed.
  

  Bee brood (pupae and larvae) although low in , has been found to be high in and , and a useful source of , , , and trace minerals , , , and . In addition, while bee brood was high in fat, it contained no fat soluble vitamins (such as A, D, and E) but it was a good source of most of the water-soluble including as well as vitamin C. The fat was composed mostly of and with 2.0% being .
  

  
  

  As alternative medicine

  
  

  is a branch of that uses honey bee products, including , , pollen, , and (Bee )..
  

  
  

  Stings

  
  

  The painful are mostly associated with the poison gland and the which are abdominal exocrine glands containing various chemicals. In , the Dufour's Gland mostly contains octadecanolide as well as some eicosanolide. There is also evidence of n-triscosane, n-, However, the secretions of these glands could also be used for nest construction.[134]
  

  
  See also
  
  
  
  
  Notes
  
  
  
  
  1. nests in a petrified forest in Arizona, implying that bees evolved much earlier,
     
  
  
  2. 
     
  
  
  
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