Bee

Carpenter bee, Xylocopa virginica, covered with pollen

Bees are flying insects, closely related to wasps and ants, all a part of the order hymenoptera. There are slightly fewer than 20,000 known species of bee, though many are undescribed and the actual number is probably much higher. They are found on every continent except Antarctica, in every habitat on the planet that contains flowering dicotyledons.

Introduction

Bees are adapted for feeding on nectar and pollen, the former primarily for sugar as an energy source, and the latter primarily for protein and other nutrients. Most pollen is used as food for larvae.

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 thirteen segments in males and twelve in females. 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.

The smallest bee is the dwarf bee (Trigona minima) and it is about 2.1 mm (5/64") long. The largest bee in the world is Megachile pluto, which can grow to a size of 39 mm (1.5"). The most common type of bee in the Northern Hemisphere are the many species of Halictidae, or sweat bees, though they are small and often mistaken for wasps or flies.

The most well-known bee species is the Western 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.

Pollination

Melissodes bee with a load of sunflower pollen

Honey bee with pollen basket of sunflower pollen

A bumblebee has fallen prey to a camouflaged green lynx pider

Bees play an important role in pollinating flowering plants, and are the major type of pollinators in ecosystems that contain flowering plants. Bees may focus on gathering nectar or on gathering pollen, depending on their greater need at the time, 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 this accomplished by bees.

Bees are extremely important as pollinators in agriculture, especially the domesticated Western honey bee, with contract pollination having overtaken the role of honey production for beekeepers in many countries. Monoculture and pollinator decline (of many bee species) have increasingly caused honey bee keepers to become migratory so that bees can be concentrated in areas of pollination need at the appropriate season. Recently, many such migratory beekeepers have experienced substantial losses, prompting the announcement of investigation into the phenomenon, dubbed "Colony Collapse Disorder," amidst great concern over the nature and extent of the losses.

Many other species of bees such as mason bees are increasingly cultured and used to meet the agricultural pollination need. Bees also play a major, though not always understood, role in providing food for birds and wildlife. Many of these bees survive in refuge in wild areas away from agricultural spraying, only to be poisoned in massive spray programs for mosquitoes, gypsy moths, or other insect pests.

Most bees are fuzzy and carry an electrostatic charge, thus aiding 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, but many 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. With solitary species, pollen and nectar are usually combined together 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").

Visiting flowers is a dangerous occupation with high mortality rates. Mantids, assassin bugs and crab spiders hide in flowers or foliage to capture unwary bees. Others are lost to birds in flight. Insecticides used on blooming plants can kill large numbers of 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, simply to replace daily casualties.

The pollination value of bees depends partly on the individual efficiency of the bees, but also on the population itself. Thus, while individual 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 the enormous difference in the population of a colony. Bee populations also vary seasonally. During early spring orchard blossoms, bumblebee populations are limited to only a few queens, and thus are not significant pollinators of early fruit.

Evolution

Bees, like ants, are essentially a highly specialized form of wasp. The ancestors of bees are thought to be 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 that were flower visitors and were partially covered with pollen when they were fed to the wasp larvae. This same evolutionary scenario has also occurred within the vespoid wasps, where the group known as "pollen wasps" also evolved from predatory ancestors. The oldest definitive bee fossil is Cretotrigona prisca in New Jersey amber and of Cretaceous age. The recently reported bee fossil, of the genus Melittosphex, is in fact a wasp stem-group to Anthophila but cannot be considered an actual bee as it lacks definitive bee traits and no information is available on whether or not it fed its larvae pollen.

The earliest animal-pollinated flowers were probably pollinated by 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 much more efficient at the task than beetles, flies, butterflies, pollen wasps, or any other pollinating insect. The appearance of such floral specialists is believed to have driven the adaptive radiation of the angiosperms, and, in turn, the bees themselves.

Eusocial and semisocial bees

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.

Many solitary bees species are gregarious, nesting together in the same area. In other bee species, groups of cohabiting females may be sisters, and if there is a division of labor within the group, then they are considered semisocial.

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 been rarely 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.

Highly eusocial bees live in colonies. Each colony has a single queen, together with workers and, at certain stages in the colony cycle, drones. When humans provide a home for a colony, the structure is called a hive. A honey bee hive can contain up to 40,000 bees at their annual peak, which occurs in the spring, but usually have fewer.

Bumblebees

Bumblebees (Bombus terrestris, B. 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 nest cavity (pre-existing), 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.

Stingless bees

Stingless bees are very diverse in behavior, but all are highly eusocial. They practice mass provisioning, complex nest architecture, and perennial colonies.

Honey bees

The true honey bees (genus Apis) have arguably the most complex social behavior among the bees. The Western (or European) honey bee, Apis mellifera, is the best known bee species and one of the best known of all insects.

Africanized honey bee

Africanized bees, also called killer bees, are a hybrid strain of Apis mellifera derived from experiments to cross European and African honey bees by Warwick Estevam Kerr. Several queen bees escaped his laboratory in South America and have spread throughout the Americas. Africanized honey bees are are better adapted to the tropics, but 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 (see diseases of the honey bee), but have their own unique parasites, pests and diseases.

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 few species of solitary bees are being increasingly cultured for commercial pollination.

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 that 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 creosotebush 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 for 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.

Cleptoparasitic bees

Cleptoparasitic bees, commonly called "cuckoo bees" because their behavior is similar to cuckoo birds, occur in several bee families, though the name is technically best applied to 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 if the female cleptoparasite has not already done so, kills and eats the host larva. In a few cases where the hosts are social species, the cleptoparasite remains in the host nest and lays many eggs, sometimes even killing the host queen and replacing her.

Many cleptoparasitic bees are closely related to, and resemble, their hosts in looks and size, (i.e., the Bombus subgenus Psithyrus, which are parasitic bumble bees that infiltrate nests of species in other subgenera of Bombus). This common pattern gave rise to the ecological principle known as "Emery's Rule". Others parasitize bees in different families, like Townsendiella, a nomadine apid, one species of which is a cleptoparasite of the melittid genus Hesperapis, while the other species in the same genus attack halictid bees.

"Nocturnal" 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.

Bee flight

In 1934 August Magnan, a French entomologist, and his assistant André Sainte-Lague claimed that current insight into the aerodynamics of flight was unable to account for flight in bumblebees.

In 2005, scientists at Caltech "demystified" honey bee flight with the assistance of high-speed digital photography and a giant robotic mock-up of a bee wing.

Miscellaneous

Bees figure prominently in mythology. See Bee (mythology).

Bees are the favorite meal of Merops apiaster, a bird. Other common predators are kingbirds, mockingbirds, bee wolves, and dragonflies.

Yellowjackets and hornets, especially when encountered as flying pests, are often mischaracterized as "bees".

Bees are often affected or even harmed by encounters with toxic chemicals in the environment (for example, see Bees and toxic chemicals).

Despite the honey bee's painful sting and the typical attitude towards insects as pests, people generally hold bees in high regard. This is most likely due to their usefulness as pollinators and as producers of honey, their social nature, and their diligence. Although a honey bee sting can be deadly to those with allergies, virtually all other bee species are non-aggressive if undisturbed, and many cannot sting at all. Bees are used to advertise many products, particularly honey and foods made with honey, thus being one of the few insects used on advertisements.

Bee Wilson (2004: p.4) states that a community of honey bees have often been employed historically by political theorists as a model of human society:

This image 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. (NB: original quotation not meta-enhanced.)

References

• From NPR's All Things Considered, March 29, 2007
• Wilson, Bee (2004). The Hive: The Story Of The Honeybee. London, Great Britain: John Murray (Publishers). ISBN 0 7195 6598 7
• Grimaldi, D. and Engel, M.S. (2005). Evolution of the Insects. Cambridge University Press. ISBN 0-521-82149-5. 
• The Bees of the World, C. D. Michener (2000)
• Monographia Apum Angliae, William Kirby (1802)

External links

• All Living Things Images, identification guides, and maps of bees
• Bee Genera of the World
• Bees, Wasps and Ants Recording Society (UK)
• Carl Hayden Bee Research Center
• Rescuing Australian stingless bees
• The first bee of spring
• Solitary Bees & Things Solitary Bees in British gardens
• Scientists identify the oldest known bee, a 100 million-year-old specimen preserved in amber
• Search for North American species at Bugguide here
• For Hymeoptera Bees and other related Insects Natural History of Bees Wasps and Insects
• Bee images on Morphbank, biological image database