Temporal range: Early Cretaceous – Recent, 100–0Ma
|Apis mellifera (Honeybee).|
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently considered as a clade Anthophila. There are nearly 20,000 known species of bees in seven to nine recognized families, though many are undescribed and the actual number is probably higher. They are found on every continent except Antarctica, in every habitat on the planet that contains insect-pollinated flowering plants.
Bees have a long proboscis (a complex "tongue") that enables them to obtain the nectar from flowers. They have antennae almost universally made up of 13 segments in males and 12 in females, as is typical for the superfamily. Bees all have two pairs of wings, the hind pair being the smaller of the two; in a very few species, one sex or caste has relatively short wings that make flight difficult or impossible, but none are wingless.
Tiny stingless bee species exist whose workers are less than 2 mm (0.079 in) long. The largest bee in the world is Megachile pluto, a leafcutter bee whose females can attain a length of 39 mm (1.5"). Members of the family Halictidae, or sweat bees, are the most common type of bee in the Northern Hemisphere, though they are small and often mistaken for wasps or flies.
The best-known bee species is the European honey bee, which, as its name suggests, produces honey, as do a few other types of bee. Human management of this species is known as beekeeping or apiculture.
- 1 Evolution
- 2 Eusocial and semisocial bees
- 3 Solitary and communal bees
- 4 Brood parasites
- 5 Nocturnal bees
- 6 Flight
- 7 Pollination
- 8 Bees and humans
- 9 See also
- 10 References
- 11 Further reading
- 12 External links
The ancestors of bees were wasps in the family Crabronidae, and therefore 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 also have occurred within the vespoid wasps, where the group known as "pollen wasps" also evolved from predatory ancestors. Up until recently, the oldest non-compression bee fossil had been Cretotrigona prisca in New Jersey amber and of Cretaceous age, a meliponine. A recently reported bee fossil, of the genus Melittosphex, is considered "an extinct lineage of pollen-collecting Apoidea sister to the modern bees", and dates from the early Cretaceous (~100 mya). Derived features of its morphology ("apomorphies") place it clearly within the bees, but it retains two unmodified ancestral traits ("plesiomorphies") of the legs (two mid-tibial spurs, and a slender hind basitarsus), indicative of its transitional status.
The earliest animal-pollinated flowers were pollinated by insects such as beetles, so the syndrome of insect pollination was well established before bees first appeared. The novelty is that bees are specialized as pollination agents, with behavioral and physical modifications that specifically enhance pollination, and are generally more efficient at the task than any other pollinating insect such as beetles, flies, butterflies and pollen wasps. The appearance of such floral specialists is believed to have driven the adaptive radiation of the angiosperms, and, in turn, the bees themselves.
Among living bee groups, the "short-tongued" bee family Colletidae has traditionally been considered the most "primitive", and sister taxon to the remainder of the bees. In the 21st century, however, some researchers have claimed that the Dasypodaidae is the basal group, the short, wasp-like mouthparts of colletids being the result of convergent evolution, rather than indicative of a plesiomorphic condition. This subject is still under debate, and the phylogenetic relationships among bee families are poorly understood.
Bees may be solitary or may live in various types of communities. The most advanced of these are eusocial colonies found among the honey bees, bumblebees, and stingless bees. Sociality, of several different types, is believed to have evolved separately many times within the bees.
If, in addition to a division of labor, the group consists of a mother and her daughters, then the group is called eusocial. The mother is considered the queen and the daughters are workers. These castes may be purely behavioral alternatives, in which case the system is considered "primitively eusocial" (similar to many paper wasps), and if the castes are morphologically discrete, then the system is "highly eusocial".
There are many more species of primitively eusocial bees than highly eusocial bees, but they have rarely been studied. The biology of most such species is almost completely unknown. The vast majority are in the family Halictidae, or "sweat bees". Colonies are typically small, with a dozen or fewer workers, on average. The only physical difference between queens and workers is average size, if they differ at all. Most species have a single season colony cycle, even in the tropics, and only mated females (future queens, or "gynes") hibernate (called diapause). A few species have long active seasons and attain colony sizes in the hundreds. The orchid bees include a number of primitively eusocial species with similar biology. Certain species of allodapine bees (relatives of carpenter bees) also have primitively eusocial colonies, with unusual levels of interaction between the adult bees and the developing brood. This is "progressive provisioning"; a larva's food is supplied gradually as it develops. This system is also seen in honey bees and some bumblebees.
Thanks to new molecular phylogeny data, a timeline of the evolution of eusocial bee species is available. The complexity of social behaviors seems positively correlated with age of evolutionary origin. This ranges from around 21 Mya in Halictidae, 53 Mya in allodapines, with the corbiculate Apidae (the group that includes honey bees and bumblebees) being the oldest at roughly 87 Mya.
Highly eusocial bees live in colonies. Each colony has a single queen, many workers and, at certain stages in the colony cycle, drones. When humans provide the nest, it is called a hive. Honey bee hives can contain up to 40,000 bees at their annual peak, which occurs in the spring, but usually have fewer.
Bumblebees (Bombus terrestris, Bombus pratorum, et al.) are eusocial in a manner quite similar to the eusocial Vespidae such as hornets. The queen initiates a nest on her own (unlike queens of honey bees and stingless bees which start nests via swarms in the company of a large worker force). 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 are rarely perennial. Bumblebee queens sometimes seek winter safety in honey bee hives, where they are sometimes found dead in the spring by beekeepers, presumably stung to death by the honey bees. It is unknown whether any survive winter in such an environment.
Bumblebees are one of the more important wild pollinators, but have declined significantly in recent decades. In the UK, two species have become nationally extinct during the last 75 years while others have been placed on the UK Biodiversity Action Plan as priority species in recognition of the need for conservation action. In 2006 a new charity, the Bumblebee Conservation Trust, was established to coordinate efforts to conserve remaining populations through conservation and education. In 2011, the International Union for the Conservation of Nature set up the Bumblebee Specialist Group to review the threat status of all bumblebee species world-wide using the IUCN Red List criteria.
The true honey bees (genus Apis) have arguably the most complex social behavior among the bees, is the best known bee species, and one of the best known of all insects. There are 29 subspecies of Apis mellifera, native to Europe, the Middle East, and Africa.
Africanized honey bee
Africanized bees, also called killer bees, are a hybrid strain of Apis mellifera derived from experiments by Warwick Estevam Kerr to cross European and African honey bee subspecies. Several queen bees escaped from his laboratory in South America and have spread throughout the Americas. Africanized honey bees are more defensive than European honey bees.
Solitary and communal bees
Most other bees, including familiar species of bee such as the Eastern carpenter bee (Xylocopa virginica), alfalfa leafcutter bee (Megachile rotundata), orchard mason bee (Osmia lignaria) and the hornfaced bee (Osmia cornifrons) are solitary in the sense that every female is fertile, and typically inhabits a nest she constructs herself. There are no worker bees for these species. Solitary bees typically produce neither honey nor beeswax. They are immune from acarine and Varroa mites, but have their own unique parasites, pests and diseases (see also diseases of the honey bee).
Solitary bees are important pollinators, and pollen is gathered for provisioning the nest with food for their brood. Often it is mixed with nectar to form a paste-like consistency. Some solitary bees have very advanced types of pollen-carrying structures on their bodies. A very few species of solitary bees are being increasingly cultured for commercial pollination. Most of these species belong to a distinct set of genera, namely: carpenter bees, sweat bees, mason bees, polyester bees, squash bees, dwarf carpenter bees, leafcutter bees, alkali bees, digger bees.
Solitary bees are often oligoleges, in that they only gather pollen from one or a few species/genera of plants (unlike honey bees and bumblebees which are generalists). No known bees are nectar specialists; many oligolectic bees will visit multiple plants for nectar, but there are no bees which visit only one plant for nectar while also gathering pollen from many different sources. Specialist pollinators also include bee species which gather floral oils instead of pollen, and male orchid bees, which gather aromatic compounds from orchids (one of the only cases where male bees are effective pollinators). In a very few cases only one species of bee can effectively pollinate a plant species, and some plants are endangered at least in part because their pollinator is dying off. There is, however, a pronounced tendency for oligolectic bees to be associated with common, widespread plants which are visited by multiple pollinators (e.g., there are some 40 oligoleges associated with creosote bush in the US desert southwest, and a similar pattern is seen in sunflowers, asters, mesquite, etc.)
Solitary bees create nests in hollow reeds or twigs, holes in wood, or, most commonly, in tunnels in the ground. 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. Providing nest boxes for solitary bees is increasingly popular among gardeners. 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 are gregarious, preferring to make nests near others of the same species, giving the appearance to the casual observer that they are 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 there are multiple females using that same entrance on a regular basis.
Brood parasites occur in several bee families including the apid subfamily Nomadinae. Females of these bees lack pollen collecting structures (the scopa) 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 kills and eats the host larva.
The cuckoo bees in the Bombus subgenus Psithyrus are closely related to, and resemble, their hosts in looks and size. This common pattern gave rise to the ecological principle "Emery's Rule". Others parasitize bees in different families, like Townsendiella, a nomadine apid, one species of which is a cleptoparasite of the dasypodaid genus Hesperapis, while the other species in the same genus attack halictid bees.
Four bee families (Andrenidae, Colletidae, Halictidae, and Apidae) contain some species that are crepuscular (these may be either the vespertine or matinal type). These bees have greatly enlarged ocelli, which are extremely sensitive to light and dark, though incapable of forming images. Many are pollinators of flowers that themselves are crepuscular, such as evening primroses, and some live in desert habitats where daytime temperatures are extremely high.
In Antoine Magnan's 1934 book Le vol des insectes, he wrote that he and André Sainte-Laguë had applied the equations of air resistance to insects 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". This has led to a common misconception that bees "violate aerodynamic theory", but in fact it merely confirms that bees do not engage in fixed-wing flight, and that their flight is explained by other mechanics, such as those used by helicopters.
In 1996, Charlie Ellington at Cambridge University showed that vortices created by many insects’ wings and non-linear effects were a vital source of lift; vortices and non-linear phenomena are notoriously difficult areas of hydrodynamics, which has made for slow progress in theoretical understanding of insect flight.
In 2005, Michael Dickinson and his Caltech colleagues studied honey bee flight with the assistance of high-speed cinematography and a giant robotic mock-up of a bee wing. Their analysis revealed that sufficient 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 arc, it flaps approximately 230 times per second, faster than a fruitfly (200 times per second) which is 80 times smaller.
Bees play an important role in pollinating flowering plants, and are the major type of pollinator in ecosystems that contain flowering plants. Bees either focus on gathering nectar or on gathering pollen depending on demand, especially in social species. Bees gathering nectar may accomplish pollination, but bees that are deliberately gathering pollen are more efficient pollinators. It is estimated that one third of the human food supply depends on insect pollination, most of which is accomplished by bees, especially the domesticated European honey bee. Contract pollination has overtaken the role of honey production for beekeepers in many countries. Monoculture and the massive decline of many bee species (both wild and domesticated) have increasingly caused honey bee keepers to become migratory so that bees can be concentrated in seasonally varying high-demand areas of pollination.
Most bees are fuzzy and carry an electrostatic charge, which aids in the adherence of pollen. Female bees periodically stop foraging and groom themselves to pack the pollen into the scopa, which is on the legs in most bees, and on the ventral abdomen on others, and modified into specialized pollen baskets on the legs of honey bees and their relatives. Many bees are opportunistic foragers, and will gather pollen from a variety of plants, while others are oligolectic, gathering pollen from only one or a few types of plant. A small number of plants produce nutritious floral oils rather than pollen, which are gathered and used by oligolectic bees. One small subgroup of stingless bees, called "vulture bees," is specialized to feed on carrion, and these are the only bees that do not use plant products as food. Pollen and nectar are usually combined to form a "provision mass", which is often soupy, but can be firm. It is formed into various shapes (typically spheroid), and stored in a small chamber (a "cell"), with the egg deposited on the mass. The cell is typically sealed after the egg is laid, and the adult and larva never interact directly (a system called "mass provisioning").
In New Zealand scientists discovered that three genera of native bees have evolved to open flower buds of the native mistletoe Peraxilla tetrapetala. The buds cannot open themselves but are visited by birds such as the tui and bellbird which twist the top of the ripe bud. That action releases a mechanism which causes the petals to suddenly spring open, giving access to the nectar and pollen. However, when observing the native bees in the Canterbury province in the South Island, the scientists were astonished to see the bees biting the top off the buds, then pushing with their legs, occasionally popping open the buds to allow the bees to harvest the nectar and pollen, and therefore aid in the pollination of the mistletoe which is in decline in New Zealand. Nowhere else in the world have bees demonstrated ability to open explosive bird-adapted flowers.
Visiting flowers can be a dangerous occupation. Many assassin bugs and crab spiders hide in flowers to capture unwary bees. Other bees are lost to birds in flight. Insecticides used on blooming plants kill many bees, both by direct poisoning and by contamination of their food supply. A honey bee queen may lay 2000 eggs per day during spring buildup, but she also must lay 1000 to 1500 eggs per day during the foraging season, mostly to replace daily casualties, most of which are workers dying of old age. Among solitary and primitively social bees, however, lifetime reproduction is among the lowest of all insects, as it is common for females of such species to produce fewer than 25 offspring.
The population value of bees depends partly on the individual efficiency of the bees, but also on the population itself. Thus while bumblebees have been found to be about ten times more efficient pollinators on cucurbits, the total efficiency of a colony of honey bees is much greater due to greater numbers. Likewise during early spring orchard blossoms, bumblebee populations are limited to only a few queens, and thus are not significant pollinators of early fruit.
Bees and humans
Bees figure prominently in mythology and folklore and have been used by political theorists as a model for human society. Journalist Bee Wilson states that the image of a community of honey bees "occurs from ancient to modern times, in Aristotle and Plato; in Virgil and Seneca; in Erasmus and Shakespeare; Tolstoy, as well as by social theorists Bernard Mandeville and Karl Marx." They are found in heraldry where they can signify industriousness as in the Manchester bee in the crest of Manchester City Council.
Despite the honey bee's painful sting and the stereotype of insects as pests, bees are generally held in high regard. This is most likely due to their usefulness as pollinators and as producers of honey, their social nature, and their reputation for diligence. Bees are one of the few insects frequently used in advertisements in a positive manner, typically for products containing honey (such as Honey Nut Cheerios).
Although a bee sting can be deadly to those with allergies, virtually all bee species are non-aggressive if undisturbed and many cannot sting at all. Humans are often a greater danger to bees, as bees can be affected or even harmed by encounters with toxic chemicals in the environment (see also bees and toxic chemicals).
From 1972 to 2006, there was a dramatic reduction in the number of feral honey bees in the US, which are now almost absent. At the same time there was a significant though somewhat gradual decline in the number of colonies maintained by beekeepers. This decline includes the cumulative losses from all factors, such as urbanization, systematic pesticide use, tracheal and Varroa mites, and commercial beekeepers' retiring and going out of business. However, in late 2006 and early 2007 the rate of attrition reached new proportions, and the term colony collapse disorder was coined to describe the sudden disappearances.
After several years of research and concern, a team of scientists headed by Jerry Bromenshenk published a paper in October 2010 saying that a new DNA-based virus, invertebrate iridescent virus or IIV6, and the fungus Nosema ceranae were found in every killed colony the group studied. In their study they found that neither agent alone seemed deadly, but a combination of the virus and Nosema ceraneae was always 100% fatal. Bromenshenk said it is not yet clear whether one condition weakens the bees enough to be finished off by the second, or whether they somehow compound the other’s destructive power. "They're co-factors, that's all we can say at the moment. They're both present in all these collapsed colonies." Investigations into the phenomenon had occurred amidst great concern over the nature and extent of the losses. In 2009 some reports from the US suggested that 1/3 of the honey bee colonies did not survive the winter, though normal winter losses are known to be around 25%. At the end of May 2012, the Swiss government reported that about half of the bee population had not survived the winter. The main cause of the decline was thought to be the parasite varroa.
Apart from colony collapse disorder, many of the losses outside the US have also been attributed to other causes. Pesticides used to treat seeds, such as Clothianidin, Imidacloprid and Thiamethoxam, have been considered prime suspects. Other species of bees such as mason bees are increasingly cultured and used to meet the agricultural pollination need.
Native pollinators include bumblebees and solitary bees, which often survive in refuges in wild areas away from agricultural spraying, but may still be poisoned in massive spray programs for mosquitoes, gypsy moths, or other insect pests. Although pesticide use remains a concern, the major problem for wild pollinator populations is the loss of the flower-rich habitat on which they depend for food. Throughout the northern hemisphere, the last 70 or so years have seen an intensification of agricultural systems, which has decreased the abundance and diversity of wild flowers.
Legislation such as the UK's Bees Act 1980 is designed to stop the decline of bees. In April 2013 the European Union announced plans to restrict the use of certain pesticides to stop bee populations from declining further.
In 2014 the Intergovernmental Panel on Climate Change report warned that bees and other pollinators faced increased risk of extinction because of global warming. This was due to their natural habitats moving and the alteration in the seasonal behaviour of species. Climate change was causing the natural foraging areas for bees to change location but the bees were not always able to move to these new areas. Bees were also emerging at different times in the year due to global warming when flowering plants were not available.
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- Honey bee life cycle
- Insect hotel#Solitary bees and wasps
- International Union for the Study of Social Insects
- Pesticide toxicity to bees
- Proboscis extension reflex
- Schmidt Sting Pain Index
- Starr sting pain scale
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- All Living Things Images, identification guides, and maps of bees
- Bee Genera of the World
- Scientists identify the oldest known bee, a 100 million-year-old specimen preserved in amber
- North American species of bees at Bugguide
- Native Bees of North America