Bumblebee
Overview
A bumble bee is any member of the bee
Bee
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 classified by the unranked taxon name Anthophila...

 genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 Bombus, in the family Apidae
Apidae
The Apidae are a large family of bees, comprising the common honey bees, stingless bees , carpenter bees, orchid bees, cuckoo bees, bumblebees, and various other less well-known groups...

. There are over 250 known species, existing primarily in the Northern Hemisphere
Northern Hemisphere
The Northern Hemisphere is the half of a planet that is north of its equator—the word hemisphere literally means “half sphere”. It is also that half of the celestial sphere north of the celestial equator...

 although they are common in New Zealand
New Zealand
New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...

 and in the Australian state of Tasmania
Tasmania
Tasmania is an Australian island and state. It is south of the continent, separated by Bass Strait. The state includes the island of Tasmania—the 26th largest island in the world—and the surrounding islands. The state has a population of 507,626 , of whom almost half reside in the greater Hobart...

.

Bumble bees are social
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....

 insects that are characterised by black and yellow body hairs, often in bands. However, some species have orange or red on their bodies, or may be entirely black.
Encyclopedia
A bumble bee is any member of the bee
Bee
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 classified by the unranked taxon name Anthophila...

 genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 Bombus, in the family Apidae
Apidae
The Apidae are a large family of bees, comprising the common honey bees, stingless bees , carpenter bees, orchid bees, cuckoo bees, bumblebees, and various other less well-known groups...

. There are over 250 known species, existing primarily in the Northern Hemisphere
Northern Hemisphere
The Northern Hemisphere is the half of a planet that is north of its equator—the word hemisphere literally means “half sphere”. It is also that half of the celestial sphere north of the celestial equator...

 although they are common in New Zealand
New Zealand
New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...

 and in the Australian state of Tasmania
Tasmania
Tasmania is an Australian island and state. It is south of the continent, separated by Bass Strait. The state includes the island of Tasmania—the 26th largest island in the world—and the surrounding islands. The state has a population of 507,626 , of whom almost half reside in the greater Hobart...

.

Bumble bees are social
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....

 insects that are characterised by black and yellow body hairs, often in bands. However, some species have orange or red on their bodies, or may be entirely black. Another obvious (but not unique) characteristic is the soft nature of the hair (long, branched seta
Seta
Seta is a biological term derived from the Latin word for "bristle". It refers to a number of different bristle- or hair-like structures on living organisms.-Animal setae:In zoology, most "setae" occur in invertebrates....

e), called pile, that covers their entire body, making them appear and feel fuzzy. They are best distinguished from similarly large, fuzzy bees by the form of the female hind leg, which is modified to form a corbicula
Pollen basket
The pollen basket or corbicula is part of the tibia on the hind legs of the four related lineages of apid bees that used to comprise the family Apidae: the honey bees, bumblebees, stingless bees, and orchid bees...

: a shiny concave surface that is bare, but surrounded by a fringe of hairs used to transport pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

 (in similar bees, the hind leg is completely hairy, and pollen grains are wedged into the hairs for transport).

Like their relatives the honey bee
Honey bee
Honey bees are a subset of bees in the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis...

s, bumble bees feed on nectar and gather pollen to feed their young.

Biology

The blood or hemolymph
Hemolymph
Hemolymph, or haemolymph, is a fluid in the circulatory system of some arthropods and is analogous to the fluids and cells making up both blood and interstitial fluid in vertebrates such as birds and mammals...

, as in other arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

s, is carried in an open circulatory system. The body organs, "heart" (dorsal aorta), muscles, etc. are surrounded in a reservoir of blood. The dorsal aorta does pulse blood through its long tube, though, so there is a circulation of sorts.

In fertilised queens the ovaries are activated when the queen lays her egg. It passes along the oviduct
Oviduct
In non-mammalian vertebrates, the passageway from the ovaries to the outside of the body is known as the oviduct. The eggs travel along the oviduct. These eggs will either be fertilized by sperm to become a zygote, or will degenerate in the body...

 to the vagina. In the vagina there is a container called the spermatheca
Spermatheca
The spermatheca , also called receptaculum seminis , is an organ of the female reproductive tract in insects, some molluscs, oligochaeta worms and certain other invertebrates and vertebrates...

. This is where the queen stores sperm from her mating. Before she lays the egg, she will decide whether to use sperm from the spermatheca to fertilise it or not. Non-fertilised eggs grow into males, and only fertilised eggs grow into females and queens.

As in all animals, hormone
Hormone
A hormone is a chemical released by a cell or a gland in one part of the body that sends out messages that affect cells in other parts of the organism. Only a small amount of hormone is required to alter cell metabolism. In essence, it is a chemical messenger that transports a signal from one...

s play a big role in the growth and development of the bumble bee. The hormones that stimulate the development of the ovaries are suppressed in the other female worker bees while the queen remains dominant. Salivary glands in the head secrete saliva
Saliva
Saliva , referred to in various contexts as spit, spittle, drivel, drool, or slobber, is the watery substance produced in the mouths of humans and most other animals. Saliva is a component of oral fluid. In mammals, saliva is produced in and secreted from the three pairs of major salivary glands,...

, which mixes with the nectar and pollen. Saliva is also mixed into the nest materials to soften them. The fat body is a nutritional store; before hibernation, queens eat as much as they can to enlarge their fat body, and the fat in the cells is used up during hibernation.
Like all bee tongues, the bumblebee tongue (the proboscis) is a long hairy structure that extends from a sheath-like modified maxilla. The primary action of the tongue is lapping, i.e. repeated dipping of the tongue into liquid. During lapping, nectar is drawn up the proboscis by capillary action. When at rest or flying, the proboscis is kept folded under the head. The abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...

 is divided into dorsal tergites
Tergum
A tergum is the dorsal portion of an arthropod segment other than the head. The anterior edge is called the base and posterior edge is called the apex or margin. A given tergum may be divided into hardened plates or sclerites commonly referred to as tergites...

 and ventral sternites
Sternum (arthropod)
The sternum is the ventral portion of a segment of an arthropod thorax or abdomen.In insects, the sterna are usually single, large sclerites, and external...

. Wax is secreted from gland
Gland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...

s on the sternites.

The brightly coloured pile
Pile (heraldry)
In heraldry, a pile is a charge usually counted as one of the ordinaries ....

 of the bumble bee is a form of aposematic
Aposematism
Aposematism , perhaps most commonly known in the context of warning colouration, describes a family of antipredator adaptations where a warning signal is associated with the unprofitability of a prey item to potential predators...

 signal. Depending on the species and morph
Polymorphism (biology)
Polymorphism in biology occurs when two or more clearly different phenotypes exist in the same population of a species — in other words, the occurrence of more than one form or morph...

, these colours can range from entirely black, to bright yellow, red, orange, white, and pink. Thick pile can also act as insulation to keep the bee warm in cold weather. Further, when flying a bee builds up an electrostatic charge, and as flowers are usually well grounded, pollen is attracted to the bee's pile when it lands. When a pollen-covered bee enters a flower, the charged pollen is preferentially attracted to the stigma
Gynoecium
Gynoecium is most commonly used as a collective term for all carpels in a flower. A carpel is the ovule and seed producing reproductive organ in flowering plants. Carpels are derived from ovule-bearing leaves which evolved to form a closed structure containing the ovules...

 because it is better grounded than the other parts of the flower.

Bumble bees do not have ears; however, they can feel the vibrations of sounds through wood and other materials.

Habitat

Bumble bees are typically found in higher latitude
Latitude
In geography, the latitude of a location on the Earth is the angular distance of that location south or north of the Equator. The latitude is an angle, and is usually measured in degrees . The equator has a latitude of 0°, the North pole has a latitude of 90° north , and the South pole has a...

s and/or high altitude
Altitude
Altitude or height is defined based on the context in which it is used . As a general definition, altitude is a distance measurement, usually in the vertical or "up" direction, between a reference datum and a point or object. The reference datum also often varies according to the context...

s, though exceptions exist (there are a few lowland tropical
Tropics
The tropics is a region of the Earth surrounding the Equator. It is limited in latitude by the Tropic of Cancer in the northern hemisphere at approximately  N and the Tropic of Capricorn in the southern hemisphere at  S; these latitudes correspond to the axial tilt of the Earth...

 species). A few species (Bombus polaris
Bombus polaris
Bombus polaris is an Arctic bumblebee species.-Distribution:The bumblebee has a circumpolar distribution, found in Canada, arctic Alaska, Arctic islands , northern Scandinavia and Russia .-Characteristics:The bumblebee has dense fur that slows heat loss...

and B. alpinus
Bombus alpinus
Bombus alpinus is a species of bumblebee. It is found in Eurasia....

) range into very cold climates where other bees might not be found; B. polaris can be found in northern Ellesmere Island
Ellesmere Island
Ellesmere Island is part of the Qikiqtaaluk Region of the Canadian territory of Nunavut. Lying within the Canadian Arctic Archipelago, it is considered part of the Queen Elizabeth Islands, with Cape Columbia being the most northerly point of land in Canada...

—the northernmost occurrence of any eusocial insect—along with its parasite, B. hyperboreus
Bombus hyperboreus
Bombus hyperboreus is an Arctic bumblebee species with a circumpolar distribution, found in arctic Canada, Alaska, Greenland, northern Scandinavia and Russia....

. One reason for this is that bumble bees can regulate their body temperature, via solar radiation
Sunlight
Sunlight, in the broad sense, is the total frequency spectrum of electromagnetic radiation given off by the Sun. On Earth, sunlight is filtered through the Earth's atmosphere, and solar radiation is obvious as daylight when the Sun is above the horizon.When the direct solar radiation is not blocked...

, internal mechanisms of "shivering" and radiative cooling from the abdomen (called heterothermy). Other bees have similar physiology
Physiology
Physiology is the science of the function of living systems. This includes how organisms, organ systems, organs, cells, and bio-molecules carry out the chemical or physical functions that exist in a living system. The highest honor awarded in physiology is the Nobel Prize in Physiology or...

, but the mechanisms have been best studied in bumble bees.

Nests

Bumble bees form colonies
Colony (biology)
In biology, a colony reference to several individual organisms of the same species living closely together, usually for mutual benefit, such as stronger defense or the ability to attack bigger prey. Some insects live only in colonies...

, which are usually much less extensive than those of honey bees. This is due to a number of factors including the small physical size of the nest cavity, the responsibility of a single female for the initial construction and reproduction that happens within the nest, and the restriction of the colony to a single season (in most species). Often, mature bumble bee nests will hold fewer than 50 individuals. Bumble bee nests may be found within tunnels in the ground made by other animals, or in tussock grass
Tussock (grass)
Tussock grasses or bunch grasses are found as native plants in natural ecosystems, as forage in pastures, and as ornamental grasses in gardens....

 as opposed to Carpenter Bee
Carpenter bee
Carpenter bees are large, hairy bees distributed worldwide. There are some 500 species of carpenter bee in 31 subgenera...

s that burrow into wood. Bumble bees sometimes construct a wax canopy ("involucrum") over the top of their nest for protection and insulation. Bumble bees do not often preserve their nests through the winter, though some tropical species live in their nests for several years (and their colonies can grow quite large, depending on the size of the nest cavity). In temperate species, the last generation of summer includes a number of queen
Queen bee
The term queen bee is typically used to refer to an adult, mated female that lives in a honey bee colony or hive; she is usually the mother of most, if not all, the bees in the hive. The queens are developed from larvae selected by worker bees and specially fed in order to become sexually mature...

s who overwinter
Overwinter
To overwinter is to pass through or wait out the winter season, or to pass through that period of the year when “winter” conditions make normal activity or even survival difficult or near impossible...

 separately in protected spots. The queens can live up to one year, possibly longer in tropical species.

Colony cycle

Bumble bee nests are first constructed by over-wintered queens in the spring (in temperate areas). Upon emerging from hibernation
Hibernation
Hibernation is a state of inactivity and metabolic depression in animals, characterized by lower body temperature, slower breathing, and lower metabolic rate. Hibernating animals conserve food, especially during winter when food supplies are limited, tapping energy reserves, body fat, at a slow rate...

, the queen collects pollen and nectar from flowers and searches for a suitable nest site. The characteristics of the nest site vary among bumble bee species, with some species preferring to nest in underground holes and others in tussock grass or directly on the ground. Once the queen finds a site, she prepares wax pots to store food, and wax cells to lay eggs in. These eggs then hatch into larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...

e, which cause the wax cells to expand isometrically into a clump of brood cells.

To develop, these larvae must be fed both nectar for carbohydrate
Carbohydrate
A carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...

s and pollen for protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

. Bumble bees feed nectar to the larvae by chewing a small hole in the brood cell into which they regurgitate nectar. Larvae are fed pollen in one of two ways, depending on the bumble bee species. So-called "pocket-maker" bumble bees create pockets of pollen at the base of the brood-cell clump that the larvae feed themselves from. Conversely, "pollen-storers" store pollen in separate wax pots and feed it to the larvae in the same fashion as nectar. Bumble bees are incapable of trophallaxis
Trophallaxis
Trophallaxis is the transfer of food or other fluids among members of a community through mouth-to-mouth or anus-to-mouth feeding. It is most highly developed in social insects such as ants, termites, wasps and bees. The word was introduced by the entomologist William Morton Wheeler in 1918...

 (direct transfer of food from one bee to another).

With proper care, the larvae progress through four instar
Instar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...

s, becoming successively larger with each moult. At the end of the fourth instar, the larvae spin silk cocoons under the wax covering the brood cells, changing them into pupal cells. The larvae then undergo an intense period of cellular growth and differentiation and become pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...

e. These pupae then develop into adult bees, and chew their way out of the silk cocoon. When adult bumble bees first emerge from their cocoons, the hairs on their body are not yet fully pigmented and are a greyish-white colour. The bees are referred to as "callow" during this time, and they will not leave the colony for at least 24 hours. The entire process from egg to adult bee can take as long as five weeks, depending on the species and the environmental conditions.

After the emergence of the first or second group of workers, workers take over the task of foraging and the queen spends most of her time laying eggs and caring for larvae. The colony grows progressively larger and at some point will begin to produce males and new queens. The point at which this occurs varies among species and is heavily dependent on resource availability and environmental factors. Unlike the workers of more advanced social insects
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....

, bumble bee workers are not physically reproductively sterile and are able to lay haploid eggs that develop into viable male bumble bees. Only fertilised queens can lay diploid eggs that mature into workers and new queens.

Early in the colony cycle, the queen bumble bee compensates for potential reproductive competition from workers by suppressing their egg-laying by way of physical aggression and pheromonal signals. Thus, the queen will usually be the mother of all of the first males laid. Workers eventually begin to lay male eggs later in the season when the queen's ability to suppress their reproduction diminishes. The reproductive competition
Competition (biology)
Competition is an interaction between organisms or species, in which the fitness of one is lowered by the presence of another. Limited supply of at least one resource used by both is required. Competition both within and between species is an important topic in ecology, especially community ecology...

 between workers and the queen is one reason that bumble bees are considered "primitively eusocial
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....

".

New queens and males leave the colony after maturation. Males in particular are forcibly driven out by the workers. Away from the colony, the new queens and males live off nectar and pollen and spend the night on flowers or in holes. The queens are eventually mated (often more than once) and search for a suitable location for diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...

 (dormancy).

Foraging behaviour

Bumble bees generally visit flowers exhibiting the bee pollination syndrome
Pollination syndrome
Pollination syndromes are suites of flower traits that have evolved in response to natural selection imposed by different pollen vectors, which can be abiotic or biotic, such as birds, bees, flies, and so forth. These traits include flower shape, size, colour, odour, reward type and amount, nectar...

. They can visit patches of flowers up to 1–2 kilometres from their colony. Bumble bees will also tend to visit the same patches of flowers every day, as long as they continue to find nectar and pollen, a habit known as pollinator or flower constancy
Flower constancy
Flower constancy or pollinator constancy is defined as the tendency of individual pollinators to exclusively visit certain flower species or morphs within a species, bypassing other available flower species that could potentially be more rewarding...

. While foraging, bumblebees can reach ground speeds of up to 15 metres per second (54 km/h).

Experiments have shown that bumble bees are able to use a combination of colour and spatial relationships in learning which flowers to forage from. After arriving at a flower, they extract nectar using their long tongue ("glossa
Glossa
Glossa may refer to several things:*glossa , a Greek word meaning "tongue" or "language" and is used in several English words including gloss, glossary, glossitis, and others...

") and store it in their crop
Crop (anatomy)
A crop is a thin-walled expanded portion of the alimentary tract used for the storage of food prior to digestion that is found in many animals, including gastropods, earthworms, leeches, insects, birds, and even some dinosaurs.- Bees :Cropping is used by bees to temporarily store nectar of flowers...

. Many species of bumble bee also exhibit what is known as "nectar robbing": instead of inserting the mouthparts into the flower normally, these bees bite directly through the base of the corolla to extract nectar, avoiding pollen transfer. These bees obtain pollen from other species of flowers that they “legitimately” visit.

Pollen is removed from flowers deliberately or incidentally by bumble bees. Incidental removal occurs when bumble bees come in contact with the anthers
Stamen
The stamen is the pollen producing reproductive organ of a flower...

 of a flower while collecting nectar. The bumble bee's body hairs receive a dusting of pollen from the anthers, which is then groomed into the corbicula
Pollen basket
The pollen basket or corbicula is part of the tibia on the hind legs of the four related lineages of apid bees that used to comprise the family Apidae: the honey bees, bumblebees, stingless bees, and orchid bees...

 ("pollen basket"). Bumblebees are also capable of buzz pollination
Buzz pollination
Sonication or buzz pollination is a technique used by some bees to release pollen which is more or less firmly held by the anthers, which makes pollination more efficient. The anther of buzz-pollinated species of plants is typically tubular, with an opening at only one end, and the pollen is inside...

.

In at least a few species, once a bumble bee has visited a flower, it leaves a scent mark on the flower. This scent mark deters visitation of the flower by other bumble bees until the scent degrades. It has been shown that this scent mark is a general chemical bouquet that bumblebees leave behind in different locations (e.g. nest, neutral, and food sites), and they learn to use this bouquet to identify both rewarding and unrewarding flowers. In addition, bumble bees rely on this chemical bouquet more when the flower has a high handling time (i.e. it takes a longer time for the bee to find the nectar).

Once they have collected nectar and pollen, bumble bees return to the nest and deposit the harvested nectar and pollen into brood cells, or into wax
Wax
thumb|right|[[Cetyl palmitate]], a typical wax ester.Wax refers to a class of chemical compounds that are plastic near ambient temperatures. Characteristically, they melt above 45 °C to give a low viscosity liquid. Waxes are insoluble in water but soluble in organic, nonpolar solvents...

 cells for storage. Unlike honey bees, bumble bees only store a few days' worth of food and so are much more vulnerable to food shortages.

Cuckoo bumblebees

Bumble bees of the subgenus Psithyrus
Psithyrus
Cuckoo bumblebees are members of the subgenus Psithyrus in the bumblebee genus Bombus. Up until recently, the 29 species of Psithyrus were considered to constitute a separate genus. They are a specialized lineage which has lost social behavior, and lost the ability to collect pollen, and are...

(known as cuckoo bumble bees, and formerly considered a separate genus) are a lineage that live parasitically
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...

 in the colonies of other bumble bees and have lost the ability to collect pollen. Before finding and invading a host colony, a Psithyrus female (there is no caste system in these species) will feed directly from flowers. Once she has infiltrated a host colony, the Psithyrus female will kill or subdue the queen of that colony and forcibly (using pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...

s and/or physical attacks) "enslave" the workers of that colony to feed her and her young. The female Psithyrus also has a number of morphological adaptations, such as larger mandibles and a larger venom sac that increase her chances of taking over a nest. Upon hatching, the male and female Psithyrus disperse and mate. Like non-parasitic bumble bee queens, female Psithyrus find suitable locations to spend the winter and enter diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...

 upon being mated.

Reproduction

In temperate zone species, in the autumn, young queens ("gyne
Gyne
Gyne is the primary reproductive female caste of social insects . Gynes are those destined to become queens, whereas female workers are typically sterile and cannot become queens...

s") mate
Mating
In biology, mating is the pairing of opposite-sex or hermaphroditic organisms for copulation. In social animals, it also includes the raising of their offspring. Copulation is the union of the sex organs of two sexually reproducing animals for insemination and subsequent internal fertilization...

 with males (drone
Drone (bee)
Drones are male honey bees. They develop from eggs that have not been fertilized, and they cannot sting, since the worker bee's stinger is a modified ovipositor .-Etymology:...

s) and diapause
Diapause
Diapause is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions...

 during the winter in a sheltered area, whether in the ground or in a man-made structure.
In the early spring, the queen comes out of diapause and finds a suitable place to create her colony. Then she builds wax
Wax
thumb|right|[[Cetyl palmitate]], a typical wax ester.Wax refers to a class of chemical compounds that are plastic near ambient temperatures. Characteristically, they melt above 45 °C to give a low viscosity liquid. Waxes are insoluble in water but soluble in organic, nonpolar solvents...

 cells in which to lay her fertilised eggs from the previous winter. The eggs that hatch develop into female workers, and in time the queen populates the colony, with workers feeding the young and performing other duties similar to honey bee workers. New reproductives are produced in autumn, and the queen and workers die, as do the males.

Sting

Queen and worker bumble bees can sting
Bee sting
A bee sting is strictly a sting from a bee . In the vernacular it can mean a sting of a bee, wasp, hornet, or yellow jacket. Some people may even call the bite of a horse-fly a bee sting...

. Unlike a honey bee
Honey bee
Honey bees are a subset of bees in the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests out of wax. Honey bees are the only extant members of the tribe Apini, all in the genus Apis...

's stinger a bumble bee's stinger lacks barbs, so they can sting more than once. Bumblebee species are not normally aggressive, but will sting in defence of their nest, or if harmed. Female cuckoo bumble bees will aggressively attack host colony members, and sting the host queen, but will ignore other animals and humans unless disturbed.

Bumble bees and people

Bumble bees are important pollinator
Pollinator
A pollinator is the biotic agent that moves pollen from the male anthers of a flower to the female stigma of a flower to accomplish fertilization or syngamy of the female gamete in the ovule of the flower by the male gamete from the pollen grain...

s of both crop
Crop
Crop may refer to:* Crop, a plant grown and harvested for agricultural use* Crop , part of the alimentary tract of some animals* Crop , a modified whip used in horseback riding or disciplining humans...

s and wildflower
Wildflower
A wildflower is a flower that grows wild, meaning it was not intentionally seeded or planted. Yet "wildflower" meadows of a few mixed species are sold in seed packets. The term "wildflower" has been made vague by commercial seedsmen who are interested in selling more flowers or seeds more...

s.

Comments by Charles Darwin

In his first (1859) edition of On the Origin of Species, Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

 wrote of "humble-bees" (a now-disused term for bumble bees; see the etymology section below in this article for more information) and their interactions with other species:
plants and animals, most remote in the scale of nature, are bound together by a web of complex relations
Ecosystem
An ecosystem is a biological environment consisting of all the organisms living in a particular area, as well as all the nonliving , physical components of the environment with which the organisms interact, such as air, soil, water and sunlight....

. [...] I have [...] reason to believe that humble-bees are indispensable to the fertilisation of the heartsease (
Viola tricolor), for other bees do not visit this flower. From experiments which I have tried, I have found that the visits of bees, if not indispensable, are at least highly beneficial to the fertilisation of our clovers; but humble-bees alone visit the common red clover (Trifolium pratense), as other bees cannot reach the nectar. Hence I have very little doubt, that if the whole genus of humble-bees became extinct or very rare in England, the heartsease and red clover would become very rare, or wholly disappear. The number of humble-bees in any district depends in a great degree on the number of field-mice
Apodemus
Apodemus is the genus of Muridae which contains the Eurasian field mice. Related to the Ryūkyū spiny rats and the prehistoric Rhagamys – and far more distantly to Mus and Malacomys –, it includes the following species:*Striped Field Mouse, Apodemus agrarius*Alpine Field Mouse, Apodemus...

, which destroy their combs and nests; and Mr. H. Newman, who has long attended to the habits of humble-bees, believes that 'more than two thirds of them are thus destroyed all over England.' Now the number of mice is largely dependent, as every one knows, on the number of cat
Cat
The cat , also known as the domestic cat or housecat to distinguish it from other felids and felines, is a small, usually furry, domesticated, carnivorous mammal that is valued by humans for its companionship and for its ability to hunt vermin and household pests...

s; and Mr. Newman says, 'Near villages and small towns I have found the nests of humble-bees more numerous than elsewhere, which I attribute to the number of cats that destroy the mice.' Hence it is quite credible that the presence of a feline animal in large numbers in a district might determine, through the intervention first of mice and then of bees, the frequency of certain flowers in that district!

Agricultural use

Bumble bees are increasingly cultured for agricultural use as pollinators because they can pollinate plant species that other pollinators cannot by using a technique known as buzz pollination. For example, bumble bee colonies are often placed in greenhouse
Greenhouse
A greenhouse is a building in which plants are grown. These structures range in size from small sheds to very large buildings...

 tomato
Tomato
The word "tomato" may refer to the plant or the edible, typically red, fruit which it bears. Originating in South America, the tomato was spread around the world following the Spanish colonization of the Americas, and its many varieties are now widely grown, often in greenhouses in cooler...

 production, because the frequency of buzzing that a bumble bee exhibits effectively releases tomato pollen.

The agricultural use of bumble bees is limited to pollination
Pollination
Pollination is the process by which pollen is transferred in plants, thereby enabling fertilisation and sexual reproduction. Pollen grains transport the male gametes to where the female gamete are contained within the carpel; in gymnosperms the pollen is directly applied to the ovule itself...

. Because bumble bees do not overwinter the entire colony, they are not obliged to stockpile honey, and are therefore not useful as honey producers.

Endangered status

Bumble bees are in danger in many developed countries due to habitat destruction
Habitat destruction
Habitat destruction is the process in which natural habitat is rendered functionally unable to support the species present. In this process, the organisms that previously used the site are displaced or destroyed, reducing biodiversity. Habitat destruction by human activity mainly for the purpose of...

 and collateral pesticide
Pesticide
Pesticides are substances or mixture of substances intended for preventing, destroying, repelling or mitigating any pest.A pesticide may be a chemical unicycle, biological agent , antimicrobial, disinfectant or device used against any pest...

 damage. In Britain
United Kingdom
The United Kingdom of Great Britain and Northern IrelandIn the United Kingdom and Dependencies, other languages have been officially recognised as legitimate autochthonous languages under the European Charter for Regional or Minority Languages...

, until relatively recently, 19 species of native true bumble bee were recognised along with six species of cuckoo bumble bees. Of these, three have been extirpated, eight are in serious decline
Pollinator decline
The term pollinator decline refers to the reduction in abundance of pollinators in many ecosystems worldwide during the end of the twentieth century....

, and only six remain widespread. Similar declines in bumble bees have been reported in Ireland
Ireland
Ireland is an island to the northwest of continental Europe. It is the third-largest island in Europe and the twentieth-largest island on Earth...

, with 4 species being designated endangered
Endangered species
An endangered species is a population of organisms which is at risk of becoming extinct because it is either few in numbers, or threatened by changing environmental or predation parameters...

, and another two species considered vulnerable
Vulnerable species
On 30 January 2010, the IUCN Red List of Threatened Species identified 9694 Vulnerable species, subspecies and varieties, stocks and sub-populations.-References:...

 to extinction. A decline in bumble bee numbers could cause large-scale changes to the countryside, leading to inadequate pollination of certain plants. The world's first bumble bee sanctuary was established at Vane Farm in the Loch Leven National Nature Reserve
Loch Leven
Loch Leven is a fresh water loch in Perth and Kinross council area, central Scotland.Roughly triangular, the loch is about 6 km at its longest. The burgh of Kinross lies at its western end. Loch Leven Castle lies on an island a short way offshore...

 in Scotland in 2008.

Some bumble bees native to North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

 are also vanishing, such as Bombus terricola, Bombus affinis and Bombus occidentalis
Bombus occidentalis
Bombus occidentalis is commonly known as the Western Bumble Bee. Bombus occidentalis is just one of the approximately thirty bumble bee species currently present in the western United States and western Canada...

, with one, Bombus franklini, that may even be extinct.

Flight

According to 20th century folklore
Folklore
Folklore consists of legends, music, oral history, proverbs, jokes, popular beliefs, fairy tales and customs that are the traditions of a culture, subculture, or group. It is also the set of practices through which those expressive genres are shared. The study of folklore is sometimes called...

, the laws of aerodynamics
Aerodynamics
Aerodynamics is a branch of dynamics concerned with studying the motion of air, particularly when it interacts with a moving object. Aerodynamics is a subfield of fluid dynamics and gas dynamics, with much theory shared between them. Aerodynamics is often used synonymously with gas dynamics, with...

 prove that the bumble bee should be incapable of flight
Flight
Flight is the process by which an object moves either through an atmosphere or beyond it by generating lift or propulsive thrust, or aerostatically using buoyancy, or by simple ballistic movement....

, as it does not have the capacity (in terms of wing size or beats per second) to achieve flight with the degree of wing loading
Wing loading
In aerodynamics, wing loading is the loaded weight of the aircraft divided by the area of the wing. The faster an aircraft flies, the more lift is produced by each unit area of wing, so a smaller wing can carry the same weight in level flight, operating at a higher wing loading. Correspondingly,...

 necessary. The origin of this claim has been difficult to pin down with any certainty. John McMasters
John H. McMasters
John H. McMasters was an aeronautical engineer notable for his contributions to aerodynamics and engineering education.-Career:...

 recounted an anecdote about an unnamed Swiss aerodynamicist at a dinner party who performed some rough calculations and concluded, presumably in jest, that according to the equations, bumble bees cannot fly. In later years McMasters has backed away from this origin, suggesting that there could be multiple sources, and that the earliest he has found was a reference in the 1934 French book ; they had applied the equations of air resistance
Drag (physics)
In fluid dynamics, drag refers to forces which act on a solid object in the direction of the relative fluid flow velocity...

 to insects and found that their flight was impossible, but that "One shouldn't be surprised that the results of the calculations don't square with reality".

Some credit physicist Ludwig Prandtl (1875–1953) of the University of Göttingen in Germany with popularizing the idea. Others say it was Swiss gas dynamicist Jacob Ackeret (1898–1981) who did the calculations.

In 1934, French entomologist Antoine Magnan included the following passage in the introduction to his book :





This translates to:


First prompted by what is done in aviation, I applied the laws of air resistance to insects, and I arrived, with Mr. Sainte-Laguë, at this conclusion that their flight is impossible.


Magnan refers to his assistant André Sainte-Laguë
André Sainte-Laguë
André Sainte-Laguë was a French mathematician who was a pioneer in the area of graph theory.His research on seat allocation methods led to one being named after him, the Sainte-Laguë method...

, a mathematician.

The calculations that purported to show that bumble bees cannot fly are based upon a simplified linear treatment of oscillating aerofoils
Airfoil
An airfoil or aerofoil is the shape of a wing or blade or sail as seen in cross-section....

. The method assumes small amplitude oscillations without flow separation. This ignores the effect of dynamic stall, an airflow separation inducing a large vortex
Vortex
A vortex is a spinning, often turbulent,flow of fluid. Any spiral motion with closed streamlines is vortex flow. The motion of the fluid swirling rapidly around a center is called a vortex...

 above the wing, which briefly produces several times the lift of the aerofoil in regular flight. More sophisticated aerodynamic analysis shows that the bumblebee can fly because its wings encounter dynamic stall in every oscillation cycle
Oscillation
Oscillation is the repetitive variation, typically in time, of some measure about a central value or between two or more different states. Familiar examples include a swinging pendulum and AC power. The term vibration is sometimes used more narrowly to mean a mechanical oscillation but sometimes...

.

Additionally, John Maynard Smith
John Maynard Smith
John Maynard Smith,His surname was Maynard Smith, not Smith, nor was it hyphenated. F.R.S. was a British theoretical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics under the well-known biologist J.B.S....

, a noted biologist with a strong background in aeronautics, has pointed out that bumble bees would not be expected to sustain flight, as they would need to generate too much power given their tiny wing area. However, in aerodynamics experiments with other insects he found that viscosity at the scale of small insects meant that even their small wings can move a very large volume of air relative to the size, and this reduces the power required to sustain flight by an order of magnitude.

Another description of a bee's wing function is that the wings work similarly to helicopter
Helicopter
A helicopter is a type of rotorcraft in which lift and thrust are supplied by one or more engine-driven rotors. This allows the helicopter to take off and land vertically, to hover, and to fly forwards, backwards, and laterally...

 blades, "reverse-pitch semirotary helicopter blades".

Bees beat their wings approximately 200 times a second. Their thorax muscles do not expand and contract on each nerve firing but rather vibrate like a plucked rubber band.

Buzz

One common, yet incorrect, assumption is that the buzz
Buzz
-People:*Buzz Aldrin , American pilot and astronaut, second person to set foot on the Moon*George Beurling , Canadian World War II fighter pilot*Buzz Hargrove , National President of the Canadian Auto Workers trade union...

ing sound of bees is caused by the beating of their wings. The sound is actually the result of the bee vibrating its flight muscle
Muscle
Muscle is a contractile tissue of animals and is derived from the mesodermal layer of embryonic germ cells. Muscle cells contain contractile filaments that move past each other and change the size of the cell. They are classified as skeletal, cardiac, or smooth muscles. Their function is to...

s, and this can be achieved while the muscles are decoupled from the wings. This is especially pronounced in bumblebees, as they must warm up their bodies considerably to get airborne at low ambient temperatures. Bumble bees have been known to reach an internal thoracic temperature of 30 °C (86 °F) using this method.

Selected species

For a complete list, see List of world bumblebee species.
  • Bombus fraternus
    Bombus fraternus
    Bombus fraternus is a relatively uncommon species of bumblebee native to the United States east of the Rocky Mountains. It is most often encountered in the Southeast, in areas with sandy soil...

  • New garden bumble bee, Bombus hypnorum
    Bombus hypnorum
    Bombus hypnorum, the tree bumblebee or new garden bumblebee, is a species of bumblebee. It is one of the most common bumblebee species in north and eastern Europe, but it's less common past 70 degrees north. Its distribution ranges over most of Europe and parts of Eastern Asia, although less common...

  • Early bumblebee
    Early Bumblebee
    The early bumblebee, Bombus pratorum, is a black bumblebee, with a yellow band around the front of the thorax and red colouration on abdominal segments 4 to 6...

    , Bombus pratorum
  • Orange-belted bumble bee Bombus ternarius
  • Buff-tailed bumble bee, or large earth bumblebee, Bombus terrestris
    Bombus terrestris
    Bombus terrestris, the buff-tailed bumblebee or large earth bumblebee is one of the most numerous bumblebee species in Europe. The queen is 2–2.7 cm long, while the workers are 1½–2 cm...


Associated parasites

  • Tracheal mites – Locustacarus buchneri
    Locustacarus buchneri
    Locustacarus buchneri is a parasitic mite that lives in the respiratory air sacs of bumblebees....

  • Protozoa
    Protozoa
    Protozoa are a diverse group of single-cells eukaryotic organisms, many of which are motile. Throughout history, protozoa have been defined as single-cell protists with animal-like behavior, e.g., movement...

    ns – Crithidia bombi
    Crithidia
    Crithidia are members of the trypanosome protozoa. They are parasites that exclusively parasitise arthropods, mainly insects. They pass from host to host as cysts in infective faeces and typically, the parasites develop in the digestive tracts of insects and interact with the intestinal epithelium...

  • Microsporidia
    Microsporidia
    The microsporidia constitute a phylum of spore-forming unicellular parasites. They were once thought to be protists but are now known to be fungi. Loosely 1500 of the probably more than one million species are named now. Microsporidia are restricted to animal hosts, and all major groups of animals...

     – Nosema bombi

Etymology

According to the Oxford English Dictionary
Oxford English Dictionary
The Oxford English Dictionary , published by the Oxford University Press, is the self-styled premier dictionary of the English language. Two fully bound print editions of the OED have been published under its current name, in 1928 and 1989. The first edition was published in twelve volumes , and...

 (OED), the term bumble bee was first recorded as having been used in the English language in the 1530 work Lesclarcissement by John Palsgrave
John Palsgrave
John Palsgrave was a priest of Henry VIII of England's court. He is known as a tutor in the royal household, and as a textbook author.-Life:...

, "I bomme, as a bombyll bee dothe." However the OED also states that the term humble bee predates it, having first been used in 1450 in Fysshynge wyth Angle, "In Juyll the greshop & the humbylbee in the medow." The latter term was used in A Midsummer Night's Dream
A Midsummer Night's Dream
A Midsummer Night's Dream is a play that was written by William Shakespeare. It is believed to have been written between 1590 and 1596. It portrays the events surrounding the marriage of the Duke of Athens, Theseus, and the Queen of the Amazons, Hippolyta...

(circa 1600) by William Shakespeare
William Shakespeare
William Shakespeare was an English poet and playwright, widely regarded as the greatest writer in the English language and the world's pre-eminent dramatist. He is often called England's national poet and the "Bard of Avon"...

, "The honie-bags steale from the humble Bees." In the period prior to World War I
World War I
World War I , which was predominantly called the World War or the Great War from its occurrence until 1939, and the First World War or World War I thereafter, was a major war centred in Europe that began on 28 July 1914 and lasted until 11 November 1918...

 the preferred English common name was humble bee, as found in On the Origin of Species (1859) by Charles Darwin (see above in this article for a lengthy quotation), though bumble bee was still in use as well, for example in The Tale of Mrs. Tittlemouse
The Tale of Mrs. Tittlemouse
The Tale of Mrs. Tittlemouse is a children's book written and illustrated by Beatrix Potter, and published by Frederick Warne & Co. in 1910. The tale is about housekeeping and insect pests in the home, and reflects Potter's own sense of tidiness and her abhorrence of insect infestations. The...

(1910) by Beatrix Potter
Beatrix Potter
Helen Beatrix Potter was an English author, illustrator, natural scientist and conservationist best known for her imaginative children’s books featuring animals such as those in The Tale of Peter Rabbit which celebrated the British landscape and country life.Born into a privileged Unitarian...

, "Suddenly round a corner, she met Babbitty Bumble--"Zizz, Bizz, Bizzz!" said the bumble bee." In the post-World War II
World War II
World War II, or the Second World War , was a global conflict lasting from 1939 to 1945, involving most of the world's nations—including all of the great powers—eventually forming two opposing military alliances: the Allies and the Axis...

 era however, humble bee has fallen into near-total disuse.

Cultural references

The orchestral interlude "Flight of the Bumblebee
Flight of the Bumblebee
"Flight of the Bumblebee" is an orchestral interlude written by Nikolai Rimsky-Korsakov for his opera The Tale of Tsar Saltan, composed in 1899–1900. The piece closes Act III, Tableau 1, during which the magic Swan-Bird changes Prince Gvidon Saltanovich into an insect so that he can fly away to...

" was composed (circa 1900) by Nikolai Rimsky-Korsakov
Nikolai Rimsky-Korsakov
Nikolai Andreyevich Rimsky-Korsakov was a Russian composer, and a member of the group of composers known as The Five.The Five, also known as The Mighty Handful or The Mighty Coterie, refers to a circle of composers who met in Saint Petersburg, Russia, in the years 1856–1870: Mily Balakirev , César...

 to represent the turning of Prince Guidon to visit his father, Tsar Saltan, in the opera The Tale of Tsar Saltan
The Tale of Tsar Saltan
The Tale of Tsar Saltan, of His Son the Renowned and Mighty Bogatyr Prince Gvidon Saltanovich, and of the Beautiful Princess-Swan is an 1831 poem by Aleksandr Pushkin, written after the Russian fairy tale edited by Vladimir Dahl...

, although the music is considered to more accurately reflect the flight of a bluebottle than a bumblebee. The music inspired Walt Disney
Walt Disney
Walter Elias "Walt" Disney was an American film producer, director, screenwriter, voice actor, animator, entrepreneur, entertainer, international icon, and philanthropist, well-known for his influence in the field of entertainment during the 20th century. Along with his brother Roy O...

 to feature a bumble bee in his 1940 animated musical Fantasia
Fantasia (film)
Fantasia is a 1940 American animated film produced by Walt Disney and released by Walt Disney Productions. The third feature in the Walt Disney Animated Classics series, the film consists of eight animated segments set to pieces of classical music conducted by Leopold Stokowski, seven of which are...

and have it sound as if it were flying in all parts of the theater. This early attempt at "surround sound
Surround sound
Surround sound encompasses a range of techniques such as for enriching the sound reproduction quality of an audio source with audio channels reproduced via additional, discrete speakers. Surround sound is characterized by a listener location or sweet spot where the audio effects work best, and...

" was unsuccessful, and the music was excluded from the film's release.

The archaic English colloquialism
Colloquialism
A colloquialism is a word or phrase that is common in everyday, unconstrained conversation rather than in formal speech, academic writing, or paralinguistics. Dictionaries often display colloquial words and phrases with the abbreviation colloq. as an identifier...

 dumbledor (also used for cockchafer
Cockchafer
The cockchafer is a European beetle of the genus Melolontha, in the family Scarabaeidae....

s) is the source of the name Albus Dumbledore
Albus Dumbledore
Professor Albus Percival Wulfric Brian Dumbledore is a major character in J. K. Rowling's Harry Potter series. For most of the series, he is the headmaster of the wizarding school Hogwarts...

, a fictional character from the Harry Potter
Harry Potter
Harry Potter is a series of seven fantasy novels written by the British author J. K. Rowling. The books chronicle the adventures of the adolescent wizard Harry Potter and his best friends Ron Weasley and Hermione Granger, all of whom are students at Hogwarts School of Witchcraft and Wizardry...

series (1997-2007). J.K. Rowling said the name "seemed to suit the headmaster, because one of his passions is music and I imagined him walking around humming to himself".

The generic epithet Bombus, assigned by Pierre André Latreille
Pierre André Latreille
Pierre André Latreille was a French zoologist, specialising in arthropods. Having trained as a Roman Catholic priest before the French Revolution, Latreille was imprisoned, and only regained his freedom after recognising a rare species he found in the prison, Necrobia ruficollis...

 in 1802, is derived from the Latin word for a buzzing or humming sound.

See also

  • Apiology
    Apiology
    Apiology is the scientific study of honey bees, a subdiscipline of melittology, which is itself a branch of entomology...

  • Bee
    Bee
    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 classified by the unranked taxon name Anthophila...

    s
  • Characteristics of common wasps and bees
    Characteristics of common wasps and bees
    While easily confused at a distance or without close observation, there are many different characteristics of common large bees and wasps which can be used to identify them.-External links:* * *...

  • Imidacloprid effects on bee population
    Imidacloprid effects on bee population
    Imidacloprid is a systemic neonicotinoid insecticide produced by the German chemical firm Bayer CropScience and sold under such trade names as Gaucho, Admire, Merit, Advantage, Confidor, Provado, and Winner. It acts as a neurotoxin and interferes with the transmission of nerve impulses in insects...

  • List of world bumblebee species
  • Regent (insecticide)
    Regent (insecticide)
    Regent is a trademark for a broad spectrum systemic insecticide containing the active ingredient fipronil. Fipronil is an insecticide discovered and developed by Rhône-Poulenc between 1985-87. It was placed on the market in 1993. Regent's rights have been purchased by BASF. It acts by contact and...

  • Volucella bombylans
    Volucella bombylans
    Volucella bombylans is a large species of hoverfly.It is larger than most hoverflies at 20mm long, and looks something like a Bumble bee with a furry black, yellow and/or white body, but is given away by its head and eyes which show it is a true fly, like a Blow-fly. It occurs in several forms each...

    , a hoverfly mimic of bumble bees
  • Bumblebee Orchid
    Bumblebee Orchid
    The Bumblebee Orchid is a typical example. It has flowers that look and smell so much like female Bumblebees that males flying nearby are irresistibly drawn in by this chemical signal, stimulating them sexually. The insect gets so excited that he starts to copulate with the flower. This is termed...


Further reading

  • Michener, C.D.
    Charles Duncan Michener
    Charles Duncan Michener is an American entomologist born in Pasadena, CA.-Biography:Much of his career has been devoted to the systematics and natural history of bees. His first peer-reviewed publication was in 1934, at the age of 16. He received his B.S. in 1939 and his Ph.D. in Entomology in...

     (2000). The Bees of the World. Johns Hopkins University Press
    Johns Hopkins University Press
    The Johns Hopkins University Press is the publishing division of the Johns Hopkins University. It was founded in 1878 and holds the distinction of being the oldest continuously running university press in the United States. The Press publishes books, journals, and electronic databases...

    .
  • "Bees." World Book Encyclopedia
    World Book Encyclopedia
    The World Book Encyclopedia is an encyclopedia published in the United States. It is self-described as "the number-one selling print encyclopedia in the world." The encyclopedia is designed to cover major areas of knowledge uniformly, but it shows particular strength in scientific, technical, and...

    1998 ed.
  • Hasley, William D. "Bees." Collier's Encyclopedia
    Collier's Encyclopedia
    P.F. Collier & Son Company published Collier's New Encyclopedia from 1902–1929, initially in 16 volumes and later in 10 volumes.Collier's 11 volume National Encyclopedia replaced Collier's New Encyclopedia....

    1990 ed.
  • Freeman, Scott. Biological Science. New Jersey: Upper Saddle River, 2002.
  • Abbott, Carl, and Bartlett, John. "Bumble Bees." Encarta Encyclopedia. 2004 ed.
  • Goulson, Dave. "Bumblebees: Their Behaviour and Ecology" 2003. Oxford University Press
    Oxford University Press
    Oxford University Press is the largest university press in the world. It is a department of the University of Oxford and is governed by a group of 15 academics appointed by the Vice-Chancellor known as the Delegates of the Press. They are headed by the Secretary to the Delegates, who serves as...

    ISBN 0-19-852607-5
  • Macdonald, M. & Nisbet, G. 2006. "Highland Bumblebees: Distribution, Ecology and Conservation." HBRG, Inverness, hbrg.org.uk. ISBN 0-9552211-0-2.

External links

The source of this article is wikipedia, the free encyclopedia.  The text of this article is licensed under the GFDL.
 
x
OK