A honey bee (also spelled honeybee) is a
eusocial
Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
flying insect
The Pterygota ( grc, πτερυγωτός, pterugōtós, winged) are a subclass of insects that includes the winged insects. It also includes insect orders that are secondarily wingless (that is, insect groups whose ancestors once had wings b ...
within the
genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
''Apis'' of the
bee
Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
, all native to
Afro-Eurasia
Afro-Eurasia (also Afroeurasia, Eurafrasia or the Old World) is a landmass comprising the continents of Africa, Asia, and Europe. The terms are compound words of the names of its constituent parts. Its mainland is the largest and most populou ...
.
After bees spread naturally throughout Africa and
Eurasia
Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...
, humans became responsible for the current
cosmopolitan distribution
In biogeography, cosmopolitan distribution is the term for the range of a taxon that extends across all or most of the world in appropriate habitats. Such a taxon, usually a species, is said to exhibit cosmopolitanism or cosmopolitism. The ext ...
of honey bees, introducing multiple subspecies into
South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
(early
16th century
The 16th century begins with the Julian year 1501 ( MDI) and ends with either the Julian or the Gregorian year 1600 ( MDC) (depending on the reckoning used; the Gregorian calendar introduced a lapse of 10 days in October 1582).
The 16th cent ...
),
North America
North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
(early 17th century), and
Australia
Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
(early 19th century).
Honey bees are known for their construction of
perennial
A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
colonial
Colonial or The Colonial may refer to:
* Colonial, of, relating to, or characteristic of a colony or colony (biology)
Architecture
* American colonial architecture
* French Colonial
* Spanish Colonial architecture
Automobiles
* Colonial (1920 au ...
nests from
wax
Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to give low ...
, the large size of their colonies, and surplus production and storage of
honey
Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...
, distinguishing their hives as a prized
foraging
Foraging is searching for wild food resources. It affects an animal's Fitness (biology), fitness because it plays an important role in an animal's ability to survive and reproduce. Optimal foraging theory, Foraging theory is a branch of behaviora ...
target of many animals, including
honey badger
The honey badger (''Mellivora capensis''), also known as the ratel ( or ), is a mammal widely distributed in Africa, Southwest Asia, and the Indian subcontinent. Because of its wide range and occurrence in a variety of habitats, it is listed ...
s,
bears
Bears are carnivoran mammals of the family Ursidae. They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats throughout the North ...
and human
hunter-gatherer
A traditional hunter-gatherer or forager is a human living an ancestrally derived lifestyle in which most or all food is obtained by foraging, that is, by gathering food from local sources, especially edible wild plants but also insects, fungi, ...
s. Only eight surviving
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of honey bee are recognized, with a total of 43
subspecies
In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
, though historically 7 to 11 species are recognized. Honey bees represent only a small fraction of the roughly 20,000 known species of bees.
The best known honey bee is the
western honey bee
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
, (''Apis mellifera''), which was domesticated for honey production and crop
pollination
Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds ...
. The only other domesticated bee is the eastern honey bee (''
Apis cerana
''Apis cerana'', the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South, Southeast and East Asia. This species is the sister species of ''Apis koschevnikovi'' and both are in the same subgenus as th ...
''), which occurs in
South
South is one of the cardinal directions or Points of the compass, compass points. The direction is the opposite of north and is perpendicular to both east and west.
Etymology
The word ''south'' comes from Old English ''sūþ'', from earlier Pro ...
,
Southeast
The points of the compass are a set of horizontal, radially arrayed compass directions (or azimuths) used in navigation and cartography. A compass rose is primarily composed of four cardinal directions—north, east, south, and west—each sepa ...
, and
East Asia
East Asia is the eastern region of Asia, which is defined in both geographical and ethno-cultural terms. The modern states of East Asia include China, Japan, Mongolia, North Korea, South Korea, and Taiwan. China, North Korea, South Korea and ...
. Only members of the genus ''Apis'' are true honey bees, but some other types of bees produce and store honey, and have been kept by humans for that purpose, including the
stingless bee
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family A ...
s belonging to the genus ''
Melipona
''Melipona'' is a genus of stingless bees, widespread in warm areas of the Neotropics, from Sinaloa and Tamaulipas (México) to Tucumán and Misiones (Argentina). About 70 species are known.Grüter, C. 2020. ''Stingless Bees: Their Behaviour, Ec ...
'' and the Indian stingless or dammar bee ''
Tetragonula iridipennis
The Indian stingless bee or dammar bee, ''Tetragonula iridipennis'', is a species of bee belonging to the family Apidae, subfamily Apinae. It was first described by Frederick Smith in 1854 who found the species in what is now the island of Sri ...
''. Modern humans also use
beeswax
Beeswax (''cera alba'') is a natural wax produced by honey bees of the genus ''Apis''. The wax is formed into scales by eight wax-producing glands in the abdominal segments of worker bees, which discard it in or at the hive. The hive workers ...
in making
candle
A candle is an ignitable wick embedded in wax, or another flammable solid substance such as tallow, that provides light, and in some cases, a fragrance. A candle can also provide heat or a method of keeping time.
A person who makes candles i ...
s,
soap
Soap is a salt of a fatty acid used in a variety of cleansing and lubricating products. In a domestic setting, soaps are surfactants usually used for washing, bathing, and other types of housekeeping. In industrial settings, soaps are use ...
,
lip balm
Lip balm or lip salve is a wax-like substance applied topically to the lips to moisturize and relieve chapped or dry lips, angular cheilitis, stomatitis, or cold sores. Lip balm often contains beeswax or carnauba wax, camphor, cetyl alcohol, lan ...
s and various
cosmetics
Cosmetics are constituted mixtures of chemical compounds derived from either natural sources, or synthetically created ones. Cosmetics have various purposes. Those designed for personal care and skin care can be used to cleanse or protect ...
, as a lubricant and in mould-making using the
lost wax process
Lost-wax casting (also called "investment casting", "precision casting", or ''cire perdue'' which has been adopted into English from the French, ) is the process by which a duplicate metal sculpture (often silver, gold, brass, or bronze) i ...
.
Etymology and name
The
genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
name ''Apis'' is
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
for "bee". Although modern dictionaries may refer to ''Apis'' as either honey bee or honeybee,
entomologist
Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arach ...
Robert Snodgrass
Robert Snodgrass (born 7 September 1987) is a Scottish professional footballer who plays as a winger for Heart of Midlothian.
Snodgrass started his senior career with Livingston. He also had a loan spell at Stirling Albion before moving to E ...
asserts that
correct usage requires two words, i.e. honey bee, as it is a kind or type of bee, whereas it is incorrect to run the two words together, as in dragonfly or butterfly, because the latter are not flies,
and have no connection with dragons or butter. Honey bee, not honeybee, is the listed common name in the
Integrated Taxonomic Information System
The Integrated Taxonomic Information System (ITIS) is an American partnership of federal agencies designed to provide consistent and reliable information on the taxonomy of biological species. ITIS was originally formed in 1996 as an interagenc ...
, the
Entomological Society of America
The Entomological Society of America (ESA) was founded in 1889 and today has more than 7,000 members, including educators, extension personnel, consultants, students, researchers, and scientists from agricultural departments, health agencies, ...
Common Names of Insects Database, and the
Tree of Life Web Project
The Tree of Life Web Project is an Internet project providing information about the diversity and phylogeny of life on Earth.
This collaborative peer reviewed project began in 1995, and is written by biologists from around the world. The site h ...
.
Origin, systematics, and distribution
![HoneyBeeAnatomy](https://upload.wikimedia.org/wikipedia/commons/4/4b/HoneyBeeAnatomy.svg)
Honey bees appear to have their center of origin in
South
South is one of the cardinal directions or Points of the compass, compass points. The direction is the opposite of north and is perpendicular to both east and west.
Etymology
The word ''south'' comes from Old English ''sūþ'', from earlier Pro ...
and
Southeast Asia
Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
(including the
Philippines
The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no),
* bik, Republika kan Filipinas
* ceb, Republika sa Pilipinas
* cbk, República de Filipinas
* hil, Republ ...
), as all the extant species except ''
Apis mellifera
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
'' are native to that region. Notably, living representatives of the earliest lineages to diverge (''
Apis florea
The dwarf honey bee (or red dwarf honey bee), ''Apis florea'', is one of two species of small, wild honey bees of southern and southeastern Asia. It has a much wider distribution than its sister species, ''Apis andreniformis''. First identified i ...
'' and ''
Apis andreniformis
''Apis andreniformis'', or the black dwarf honey bee, is a relatively rare species of honey bee whose native habitat is the tropical and subtropical regions of Southeast Asia.
''A. andreniformis'' was the fifth honey bee species to be described ...
'') have their center of origin there.
The first ''Apis'' bees appear in the
fossil record
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
at the
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
-
Oligocene
The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
boundary (34
mya
Mya may refer to:
Brands and product names
* Mya (program), an intelligent personal assistant created by Motorola
* Mya (TV channel), an Italian Television channel
* Midwest Young Artists, a comprehensive youth music program
Codes
* Burmese ...
), in European deposits. The origin of these prehistoric honey bees does not necessarily indicate Europe as the place of origin of the genus, only that the bees were present in Europe by that time. Few fossil deposits are known from South Asia, the suspected region of honey bee origin, and fewer still have been thoroughly studied.
No ''Apis'' species existed in the New World during human times before the introduction of ''A. mellifera'' by Europeans. Only one fossil species is documented from the New World, ''
Apis nearctica
''Apis nearctica'' is an extinct species of honey bee which existed in what is now Nevada during the Middle Miocene period. It was discovered at Stewart Valley, Nevada, and described by Michael S. Engel, Ismael A. Hinojosa-Díaz, and Alexandr P ...
'', known from a single 14 million-year-old specimen from Nevada.
The close relatives of modern honey bees – e.g.,
bumblebee
A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
s and
stingless bee
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family A ...
s – are also social to some degree, and social behavior is considered to be a trait that predates the origin of the genus. Among the extant members of ''Apis'', the more
basal species make single, exposed combs, while the more recently evolved species nest in cavities and have multiple combs, which has greatly facilitated their domestication.
Species
While about 20,000 species of bees exist,
only eight
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of honey bee are recognized, with a total of 43
subspecies
In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
, although historically seven to 11 species are recognized:
''Apis andreniformis'' (the black dwarf honey bee); ''Apis cerana'' (the eastern honey bee); ''Apis dorsata'' (the giant honey bee); ''Apis florea'' (the red dwarf honey bee); ''
Apis koschevnikovi
''Apis koschevnikovi'', Koschevnikov's honey bee, is a species of honey bee which inhabits Malaysian and Indonesian Borneo, where it lives sympatrically with other honey bee species such as ''Apis cerana'' (specifically '' A. c. nuluensis'').
T ...
'' (Koschevnikov's honey bee); ''Apis laboriosa'' (the Himalayan giant honey bee); ''Apis mellifera'' (the western honey bee); and ''Apis nigrocincta'' (the Philippine honey bee).
Honey bees are the only extant members of the
tribe
The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...
Apini. Today's honey bees constitute three
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
s: ''Micrapis'' (the dwarf honey bees), ''Megapis'' (the giant honey bee), and ''Apis'' (the western honey bee and its close relatives).
Most species have historically been cultured or at least exploited for honey and
beeswax
Beeswax (''cera alba'') is a natural wax produced by honey bees of the genus ''Apis''. The wax is formed into scales by eight wax-producing glands in the abdominal segments of worker bees, which discard it in or at the hive. The hive workers ...
by humans indigenous to their native ranges. Only two species have been truly
domesticated
Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
: ''
Apis mellifera
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
'' and ''
Apis cerana
''Apis cerana'', the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South, Southeast and East Asia. This species is the sister species of ''Apis koschevnikovi'' and both are in the same subgenus as th ...
''. ''A. mellifera'' has been cultivated at least since the time of the building of the
Egyptian pyramid
The Egyptian pyramids are ancient masonry structures located in Egypt. Sources cite at least 118 identified "Egyptian" pyramids. Approximately 80 pyramids were built within the Kingdom of Kush, now located in the modern country of Sudan. Of ...
s, and only that species has been moved extensively beyond its native range.
''Micrapis''
''
Apis florea
The dwarf honey bee (or red dwarf honey bee), ''Apis florea'', is one of two species of small, wild honey bees of southern and southeastern Asia. It has a much wider distribution than its sister species, ''Apis andreniformis''. First identified i ...
'' and ''
Apis andreniformis
''Apis andreniformis'', or the black dwarf honey bee, is a relatively rare species of honey bee whose native habitat is the tropical and subtropical regions of Southeast Asia.
''A. andreniformis'' was the fifth honey bee species to be described ...
'' are small honey bees of southern and southeastern Asia. They make very small, exposed nests in trees and shrubs. Their stings are often incapable of penetrating human skin, so the
hive
A hive may refer to a beehive, an enclosed structure in which some honey bee species live and raise their young.
Hive or hives may also refer to:
Arts
* ''Hive'' (game), an abstract-strategy board game published in 2001
* "Hive" (song), a 201 ...
and
swarm
Swarm behaviour, or swarming, is a collective behaviour exhibited by entities, particularly animals, of similar size which aggregate together, perhaps milling about the same spot or perhaps moving ''en masse'' or migrating in some direction. ...
s can be handled with minimal protection. They occur largely
sympatric
In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...
ally, though they are very distinct
evolution
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
arily and are probably the result of
allopatric speciation
Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
, their distribution later converging.
Given that ''A. florea'' is more widely distributed and ''A. andreniformis'' is considerably more aggressive, honey is, if at all, usually harvested from the former only. They are the most ancient extant lineage of honey bees, maybe diverging in the
Bartonian
The Bartonian is, in the ICS's geologic time scale, a stage or age in the middle Eocene Epoch or Series. The Bartonian Age spans the time between . It is preceded by the Lutetian and is followed by the Priabonian Age.
Stratigraphic definitio ...
(some 40 million years ago or slightly later) from the other lineages, but do not seem to have diverged from each other a long time before the
Neogene
The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...
.
''
Apis florea
The dwarf honey bee (or red dwarf honey bee), ''Apis florea'', is one of two species of small, wild honey bees of southern and southeastern Asia. It has a much wider distribution than its sister species, ''Apis andreniformis''. First identified i ...
'' have smaller wing spans than its sister species.
''Apis florea'' are also completely yellow with the exception of the scutellum of workers, which is black.
''Megapis''
Two species are recognized in the subgenus ''Megapis''. They usually build single or a few exposed combs on high tree limbs, on cliffs, and sometimes on buildings. They can be very fierce. Periodically robbed of their honey by human "honey hunters", colonies are easily capable of
stinging
Sting may refer to:
* Stinger or sting, a structure of an animal to inject venom, or the injury produced by a stinger
* Irritating hairs or prickles of a stinging plant, or the plant itself
Fictional characters and entities
* Sting (Middle-earth ...
a human being to death if provoked.
*''
Apis dorsata
''Apis dorsata'', the giant honey bee, सिङ्गुस in Nepali, is a honey bee of South and Southeast Asia, found mainly in forested areas such as the Terai of Nepal. They are typically around long. Nests are mainly built in exposed pla ...
'', the giant honey bee, is native and widespread across most of South and Southeast Asia.
**''A. d. binghami'', the Indonesian giant honey bee, is classified as the
Indonesia
Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
n subspecies of the giant honey bee or a distinct species; in the latter case, ''A. d. breviligula'' and / or other lineages would probably also have to be considered species.
*''
Apis laboriosa
''Apis laboriosa'', the Himalayan giant honey bee, is the world’s largest honey bee; single adults can measure up to in length. Before 1980, ''Apis laboriosa'' was considered to be a subspecies of the widespread ''Apis dorsata'', the giant ho ...
'', the Himalayan giant honey bee, was initially described as a distinct species. Later, it was included in ''A. dorsata'' as a subspecies
based on the
biological species concept
The species problem is the set of questions that arises when biologists attempt to define what a species is. Such a definition is called a species concept; there are at least 26 recognized species concepts. A species concept that works well for se ...
, though authors applying a genetic species concept have suggested it should be considered a separate species
and more recent research has confirmed this classification.
[Kitnya N, Prabhudev MV, Bhatta CP, Pham TH, Nidup T, Megu K, Chakravorty J, Brockmann A, Otis GW (2020) Geographical distribution of the giant honey bee ''Apis laboriosa'' Smith, 1871 (Hymenoptera, Apidae). ZooKeys 951: 67-81. https://doi.org/10.3897/zookeys.951.49855] Essentially restricted to the
Himalayas
The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 100 ...
, it differs little from the giant honey bee in appearance, but has extensive behavioral
adaptations
In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
that enable it to nest in the open at high altitudes despite low ambient temperatures. It is the largest living honey bee.
''Apis''
![Western honey bee on a honeycomb](https://upload.wikimedia.org/wikipedia/commons/a/ae/Western_honey_bee_on_a_honeycomb.jpg)
Eastern ''Apis'' species include three or four species, including ''
A. koschevnikovi'', ''
Apis nigrocincta
''Apis nigrocincta'' is a species of honey bee that inhabits the Philippine island of Mindanao as well as the Indonesian islands of Sangihe and Sulawesi. The species is known to have queens with the highest mating frequencies of any species of ...
'', and ''
A. cerana''. The genetics of the western honey bee (''
A. mellifera'') are unclear.
Koschevnikov's honey bee
Koschevnikov's honey bee (''
Apis koschevnikovi
''Apis koschevnikovi'', Koschevnikov's honey bee, is a species of honey bee which inhabits Malaysian and Indonesian Borneo, where it lives sympatrically with other honey bee species such as ''Apis cerana'' (specifically '' A. c. nuluensis'').
T ...
'') is often referred to in the literature as the "red bee of Sabah"; however, ''A. koschevnikovi'' is pale reddish in
Sabah State
Sabah () is a state of Malaysia located in northern Borneo, in the region of East Malaysia. Sabah borders the Malaysian state of Sarawak to the southwest and the North Kalimantan province of Indonesia to the south. The Federal Territory of ...
,
Borneo
Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and eas ...
,
Malaysia
Malaysia ( ; ) is a country in Southeast Asia. The federation, federal constitutional monarchy consists of States and federal territories of Malaysia, thirteen states and three federal territories, separated by the South China Sea into two r ...
, but a dark, coppery color in the
Malay Peninsula
The Malay Peninsula (Malay: ''Semenanjung Tanah Melayu'') is a peninsula in Mainland Southeast Asia. The landmass runs approximately north–south, and at its terminus, it is the southernmost point of the Asian continental mainland. The area ...
and
Sumatra
Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
,
Indonesia
Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
.
Its habitat is limited to the tropical evergreen forests of the
Malay Peninsula
The Malay Peninsula (Malay: ''Semenanjung Tanah Melayu'') is a peninsula in Mainland Southeast Asia. The landmass runs approximately north–south, and at its terminus, it is the southernmost point of the Asian continental mainland. The area ...
,
Borneo
Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and eas ...
and
Sumatra
Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
and they do not live in tropical evergreen rain forests which extend into
Thailand
Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
,
Myanmar
Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
,
Cambodia
Cambodia (; also Kampuchea ; km, កម្ពុជា, UNGEGN: ), officially the Kingdom of Cambodia, is a country located in the southern portion of the Indochinese Peninsula in Southeast Asia, spanning an area of , bordered by Thailand t ...
and
Vietnam
Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
.
Philippine honey bee
''
Apis nigrocincta
''Apis nigrocincta'' is a species of honey bee that inhabits the Philippine island of Mindanao as well as the Indonesian islands of Sangihe and Sulawesi. The species is known to have queens with the highest mating frequencies of any species of ...
'' is a cavity-nesting species. The species has rust-colored
scapes, legs, and
clypeuses, with reddish-tan hair color that covers most of the body.
Eastern honey bee
''
Apis cerana
''Apis cerana'', the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South, Southeast and East Asia. This species is the sister species of ''Apis koschevnikovi'' and both are in the same subgenus as th ...
'', the eastern honey bee proper, is the traditional honey bee of southern and eastern Asia. One of its subspecies, the Indian honey bee (''
A. c. indica''), was
domesticated
Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
and kept in hives in a fashion similar to ''A. mellifera'', though on a more limited, regional scale.
It has not been possible yet to resolve its relationship to the Bornean honey bee ''
A. c. nuluensis'' and ''
Apis nigrocincta
''Apis nigrocincta'' is a species of honey bee that inhabits the Philippine island of Mindanao as well as the Indonesian islands of Sangihe and Sulawesi. The species is known to have queens with the highest mating frequencies of any species of ...
'' from the Philippines to satisfaction; some researchers argue that these are indeed distinct species, but that ''
A. cerana'' as defined is still
paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
, consisting of several separate species,
though other researchers argue ''cerana'' is a single monophyletic species.
Western honey bee
![Apis mellifera Tanzania](https://upload.wikimedia.org/wikipedia/commons/d/d9/Apis_mellifera_Tanzania.jpg)
''A. mellifera'', the most common domesticated species, the first domestication before 2600BC, and was the third insect whose
genome
In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
was mapped. It seems to have originated in eastern tropical
Africa
Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
and spread from there to
Europe
Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and eastwards into
Asia
Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
to the
Tian Shan
The Tian Shan,, , otk, 𐰴𐰣 𐱅𐰭𐰼𐰃, , tr, Tanrı Dağı, mn, Тэнгэр уул, , ug, تەڭرىتاغ, , , kk, Тәңіртауы / Алатау, , , ky, Теңир-Тоо / Ала-Тоо, , , uz, Tyan-Shan / Tangritog‘ ...
range. It is variously called the European, western, or common honey bee in different parts of the world. Many
subspecies
In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
have adapted to the local geographic and climatic environments; in addition, breeds such as the
Buckfast bee
The Buckfast bee is a breed of honey bee, a cross of many subspecies and their strains, developed by Brother Adam (born Karl Kehrle in 1898 in Germany), who was in charge of beekeeping from 1919 at Buckfast Abbey in Devon in the United Kingdom ...
have been bred. Behavior, color, and anatomy can be quite different from one subspecies or even strain to another.
''A. mellifera''
phylogeny
A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
is the most enigmatic of all honey bee species. It seems to have diverged from its eastern relatives only during the
Late Miocene
The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
. This would fit the hypothesis that the ancestral stock of cave-nesting honey bees was separated into the western group of East Africa and the eastern group of tropical Asia by
desertification
Desertification is a type of land degradation in drylands in which biological productivity is lost due to natural processes or induced by human activities whereby fertile areas become increasingly arid. It is the spread of arid areas caused by ...
in the
Middle East
The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
and adjacent regions, which caused declines of food plants and trees that provided nest sites, eventually causing
gene flow
In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
to cease.
The diversity of ''A. mellifera'' subspecies is probably the product of a largely
Early Pleistocene
The Early Pleistocene is an unofficial sub-epoch in the international geologic timescale in chronostratigraphy, being the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently estimated to span the time ...
radiation
In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or through a material medium. This includes:
* ''electromagnetic radiation'', such as radio waves, microwaves, infrared, visi ...
aided by climate and habitat changes during the
last ice age. That the western honey bee has been intensively managed by humans for many millennia – including hybridization and introductions – has apparently increased the speed of its
evolution
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
and confounded the DNA sequence data to a point where little of substance can be said about the exact relationships of many ''A. mellifera'' subspecies.
''Apis mellifera'' is not native to
the Americas
The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North America, North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. ...
, so it was not present when the European explorers and colonists arrived. However, other native bee species were kept and traded by indigenous peoples. In 1622, European colonists brought the
German honey bee (''A. m. mellifera'') to the Americas first, followed later by the
Italian honey bee (''A. m. ligustica'') and others. Many of the crops that depend on western honey bees for pollination have also been imported since colonial times. Escaped swarms (known as "wild" honey bees, but actually
feral
A feral () animal or plant is one that lives in the wild but is descended from domesticated individuals. As with an introduced species, the introduction of feral animals or plants to non-native regions may disrupt ecosystems and has, in some ...
) spread rapidly as far as the
Great Plains
The Great Plains (french: Grandes Plaines), sometimes simply "the Plains", is a broad expanse of flatland in North America. It is located west of the Mississippi River and east of the Rocky Mountains, much of it covered in prairie, steppe, an ...
, usually preceding the colonists. Honey bees did not naturally cross the
Rocky Mountains
The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in straight-line distance from the northernmost part of western Canada, to New Mexico in ...
; they were transported by the
Mormon
Mormons are a religious and cultural group related to Mormonism, the principal branch of the Latter Day Saint movement started by Joseph Smith in upstate New York during the 1820s. After Smith's death in 1844, the movement split into several ...
pioneers to Utah in the late 1840s, and by ship to
California
California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
in the early 1850s.
Africanized honey bee
Africanized honey bees (known colloquially as "killer bees") are
hybrid
Hybrid may refer to:
Science
* Hybrid (biology), an offspring resulting from cross-breeding
** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species
** Hybridity, the property of a hybrid plant which is a union of two dif ...
s between European stock and the East African lowland subspecies ''
A. m. scutellata''; they are often more aggressive than European honey bees and do not create as much of a honey surplus, but are more resistant to disease and are better foragers. Accidentally released from quarantine in
Brazil
Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, they have spread to North America and constitute a
pest in some regions. However, these strains do not overwinter well, so they are not often found in the colder, more northern parts of North America. The original breeding experiment for which the East African lowland honey bees were brought to Brazil in the first place has continued (though not as originally intended). Novel hybrid strains of domestic and re-domesticated Africanized honey bees combine high resilience to tropical conditions and good yields. They are popular among beekeepers in Brazil.
Living and fossil honey bees (Apini: ''Apis'')
Tribe Apini Latreille
Genus ''Apis'' Linnaeus (''sensu lato'')
*''henshawi'' species group (†''Priorapis'' Engel, †''Synapis'' Cockerell)
***†''A. vetusta'' Engel
***†''A. henshawi'' Cockerell
***†''A. petrefacta'' (Říha)
***†''A. miocenica'' Hong
***†''A. "longtibia"'' Zhang
***†''A.'' "Miocene 1"
*''armbrusteri'' species group (†''Cascapis'' Engel)
***†''A. armbrusteri'' Zeuner
***†''A. nearctica'', species novus
*''florea'' species group (''Micrapis'' Ashmead)
***''A. florea'' Fabricius
***''A. andreniformis'' Smith
*''dorsata'' species group (''Megapis'' Ashmead)
***†''A. lithohermaea'' Engel
***''A. dorsata'' Fabricius
***''A. laboriosa'' Smith
*''mellifera'' species group (''Apis'' Linnaeus ''sensu stricto'')
**''mellifera'' subgroup
***''A. mellifera'' Linnaeus (''Apis'' Linnaeus ''sensu strictissimo'')
**''cerana'' subgroup (''Sigmatapis'' Maa)
***''A. cerana'' Fabricius
***''A. nigrocincta'' Smith
***''A. koschevnikovi'' Enderlein
Life cycle
As in a few other types of
eusocial
Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
bees, a colony generally contains one
queen bee
A queen bee is typically an adult, mated female (gyne) that lives in a colony or hive of honey bees. With fully developed reproductive organs, the queen is usually the mother of most, if not all, of the bees in the beehive. Queens are developed ...
, a female; seasonally up to a few thousand
drone bee
A drone is a male honey bee. Unlike the female worker bee, drones do not have stingers. They gather neither nectar nor pollen and are unable to feed without assistance from worker bees. A drone's only role is to mate with a maiden queen in nupt ...
s, or males;
and tens of thousands of female
worker bee
A worker bee is any female (eusocial) bee that lacks the full reproductive capacity of the colony's queen bee; under most circumstances, this is correlated to an increase in certain non-reproductive activities relative to a queen. While worker be ...
s. Details vary among the different species of honey bees, but common features include:
#Eggs are laid singly in a cell in a wax
honeycomb
A honeycomb is a mass of Triangular prismatic honeycomb#Hexagonal prismatic honeycomb, hexagonal prismatic Beeswax, wax cells built by honey bees in their beehive, nests to contain their larvae and stores of honey and pollen.
beekeeping, Beekee ...
, produced and shaped by the worker bees. Using her
spermatheca
The spermatheca (pronounced plural: spermathecae ), also called receptaculum seminis (plural: receptacula seminis), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, oligochaeta worms and certain other ...
, the queen can choose to fertilize the egg she is laying, usually depending on which cell she is laying it into. Drones develop from unfertilised eggs and are
haploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, while females (queens and worker bees) develop from fertilised eggs and are
diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
. Larvae are initially fed with
royal jelly
Royal jelly is a honey bee secretion that is used in the nutrition of larvae and adult queens. It is secreted from the glands in the hypopharynx of nurse bees, and fed to all larvae in the colony, regardless of sex or caste.Graham, J. (ed.) (19 ...
produced by worker bees, later switching to honey and pollen. The exception is a larva fed solely on royal jelly, which will develop into a queen bee. The larva undergoes several moultings before spinning a cocoon within the cell, and
pupating.
#Young worker bees, sometimes called "nurse bees", clean the hive and feed the larvae. When their royal jelly-producing glands begin to
atrophy
Atrophy is the partial or complete wasting away of a part of the body. Causes of atrophy include mutations (which can destroy the gene to build up the organ), poor nourishment, poor circulation, loss of hormonal support, loss of nerve supply t ...
, they begin building comb cells. They progress to other within-colony tasks as they become older, such as receiving nectar and pollen from foragers, and guarding the hive. Later still, a worker takes her first orientation flights and finally leaves the hive and typically spends the remainder of her life as a forager.
#Worker bees cooperate to find food and use a pattern of "dancing" (known as the
bee dance
Waggle dance is a term used in beekeeping and ethology for a particular figure-eight dance of the honey bee. By performing this dance, successful foragers can share information about the direction and distance to patches of flowers yielding ne ...
or waggle dance) to communicate information regarding resources with each other; this dance varies from species to species, but all living species of ''Apis'' exhibit some form of the behavior. If the resources are very close to the hive, they may also exhibit a less specific dance commonly known as the "round dance".
#Honey bees also perform
tremble dance
A tremble dance is a dance performed by forager honey bees of the species ''Apis mellifera'' to recruit more receiver honey bees to collect nectar from the workers.
History of discovery
The tremble dance was first described by Karl von Frisch in t ...
s, which recruit receiver bees to collect nectar from returning foragers.
#Virgin queens go on mating flights away from their home colony to a drone congregation area, and mate with multiple drones before returning. The drones die in the act of mating. Queen honey bees do not mate with drones from their home colony.
#Colonies are established not by solitary queens, as in most bees, but by groups known as "
swarm
Swarm behaviour, or swarming, is a collective behaviour exhibited by entities, particularly animals, of similar size which aggregate together, perhaps milling about the same spot or perhaps moving ''en masse'' or migrating in some direction. ...
s", which consist of a mated queen and a large contingent of worker bees. This group moves ''en masse'' to a nest site which was scouted by worker bees beforehand and whose location is communicated with a special type of dance. Once the swarm arrives, they immediately construct a new wax comb and begin to raise new worker brood. This type of nest founding is not seen in any other living bee genus, though several groups of
vespid
The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
wasps also found new nests by swarming (sometimes including multiple queens). Also,
stingless bee
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family A ...
s will start new nests with large numbers of worker bees, but the nest is constructed before a queen is escorted to the site, and this worker force is not a true "swarm".
Gallery
File:Bienenwabe mit Eiern 39.jpg, Honey bee eggs shown in opened wax cells
File:Bienenwabe mit Eiern und Brut 5.jpg, Eggs and larvae
File:Drohnenpuppen 79b.jpg, Drone pupae
File:Birth of black bee (Apis mellifera mellifera).jpg, Emergence of a European dark honey bee (''A. m. mellifera'')
Winter survival
In cold climates, honey bees stop flying when the temperature drops below about and crowd into the central area of the hive to form a "winter cluster". The worker bees huddle around the queen bee at the center of the cluster, shivering to keep the center between at the start of winter (during the broodless period) and once the queen resumes laying. The worker bees rotate through the cluster from the outside to the inside so that no bee gets too cold. The outside edges of the cluster stay at about . The colder the weather is outside, the more compact the cluster becomes. During winter, they consume their stored honey to produce body heat. The amount of honey consumed during the winter is a function of winter length and severity, but ranges in temperate climates from . In addition, certain bees, including the
western honey bee
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
as well as ''
Apis cerana
''Apis cerana'', the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South, Southeast and East Asia. This species is the sister species of ''Apis koschevnikovi'' and both are in the same subgenus as th ...
'', are known to engage in effective methods of nest thermoregulation during periods of varying temperature in both summer and winter. During the summer, however, this is achieved through fanning and water evaporation from water collected in various fields.
Pollination
Of all the honey bee species, only ''A. mellifera'' has been used extensively for commercial pollination of fruit and vegetable crops. The scale of these pollination services is commonly measured in the billions of dollars, credited with adding about 9% to the value of crops across the world. However, despite contributing substantially to crop pollination, there is debate about the potential spillover to natural landscapes and competition between managed honey bees and many of the ~20,000 species of wild pollinators.
Species of ''Apis'' are generalist floral visitors, and pollinate many species of flowering plants, but because of their "generalized" nature, they often do so inefficiently. Without specialized adaptations for specific flowers, their ability to reach pollen and nectar is often limited. This combined with their behavioural flexibility may be why they are the most commonly documented
pollen thieves. Indeed, for plant species with more specialized pollinators, experiments show that increased honeybee visitation can actually reduce pollination, both where honey bees are non-native and even where they are native. What's more, their tendency to visit all species in a given area means that the pollen they carry for any one species is often very diluted. As such, they can provide some pollination to many plants, but most plants have some native pollinator that is more effective at pollinating that species. When honey bees are present as an invasive species in an area, they compete for flowers with native pollinators, which can actually push out the native species.
Claims of human dependency
Western honey bee
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
s are often described as essential to human food production, leading to claims that without their pollination humanity would starve or die out. Apples, blueberries and cherries, for example, are 90 percent dependent on honeybee pollination.
Albert Einstein
Albert Einstein ( ; ; 14 March 1879 – 18 April 1955) was a German-born theoretical physicist, widely acknowledged to be one of the greatest and most influential physicists of all time. Einstein is best known for developing the theory ...
is sometimes misquoted as saying "If bees disappeared off the face of the earth, man would only have four years left to live"''.'' Einstein did not say this and there is no science to support this prediction.
Many important crops need no insect pollination at all. The ten most important crops, comprising 60% of all human food energy, fall into this category:
plantain
Plantain may refer to:
Plants and fruits
* Cooking banana, banana cultivars in the genus ''Musa'' whose fruits are generally used in cooking
** True plantains, a group of cultivars of the genus ''Musa''
* ''Plantaginaceae'', a family of flowerin ...
s are sterile and propagated by cuttings, as are
cassava
''Manihot esculenta'', common name, commonly called cassava (), manioc, or yuca (among numerous regional names), is a woody shrub of the spurge family, Euphorbiaceae, native to South America. Although a perennial plant, cassava is extensively ...
;
potato
The potato is a starchy food, a tuber of the plant ''Solanum tuberosum'' and is a root vegetable native to the Americas. The plant is a perennial in the nightshade family Solanaceae.
Wild potato species can be found from the southern Unit ...
es,
yams, and
sweet potato
The sweet potato or sweetpotato (''Ipomoea batatas'') is a dicotyledonous plant that belongs to the Convolvulus, bindweed or morning glory family (biology), family, Convolvulaceae. Its large, starchy, sweet-tasting tuberous roots are used as a r ...
es are root vegetables propagated by
tuber
Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...
s;
soybean
The soybean, soy bean, or soya bean (''Glycine max'') is a species of legume native to East Asia, widely grown for its edible bean, which has numerous uses.
Traditional unfermented food uses of soybeans include soy milk, from which tofu an ...
s are
self-pollinated; and
rice
Rice is the seed of the grass species ''Oryza sativa'' (Asian rice) or less commonly ''Oryza glaberrima
''Oryza glaberrima'', commonly known as African rice, is one of the two domesticated rice species. It was first domesticated and grown i ...
,
wheat
Wheat is a grass widely cultivated for its seed, a cereal grain that is a worldwide staple food. The many species of wheat together make up the genus ''Triticum'' ; the most widely grown is common wheat (''T. aestivum''). The archaeologi ...
,
sorghum
''Sorghum'' () is a genus of about 25 species of flowering plants in the grass family (Poaceae). Some of these species are grown as cereals for human consumption and some in pastures for animals. One species is grown for grain, while many othe ...
, and
maize
Maize ( ; ''Zea mays'' subsp. ''mays'', from es, maíz after tnq, mahiz), also known as corn (North American and Australian English), is a cereal grain first domesticated by indigenous peoples in southern Mexico about 10,000 years ago. Th ...
are
wind-pollinated
Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Almost all gymnosperms are anemophilous, as are many plants in the order Poales, including grasses, sedges, and rushes. Other common anemophilous plan ...
, as are most other grasses.
No crops originating in the
New World
The term ''New World'' is often used to mean the majority of Earth's Western Hemisphere, specifically the Americas."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: Oxford University Press, p. 3 ...
depend on the western honey bee (''Apis mellifera'') at all, as the bee is an
invasive species
An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
brought over with colonists in the last few centuries.
Tomato
The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...
es,
pepper
Pepper or peppers may refer to:
Food and spice
* Piperaceae or the pepper family, a large family of flowering plant
** Black pepper
* ''Capsicum'' or pepper, a genus of flowering plants in the nightshade family Solanaceae
** Bell pepper
** Chili ...
s,
squash
Squash may refer to:
Sports
* Squash (sport), the high-speed racquet sport also known as squash racquets
* Squash (professional wrestling), an extremely one-sided match in professional wrestling
* Squash tennis, a game similar to squash but pla ...
, and all other New World crops evolved with native pollinators such as
squash bee
The name squash bee, also squash and gourd bee, is applied to two related genera of bees in the tribe Eucerini; '' Peponapis'' and ''Xenoglossa''. Both genera are oligoleges (pollen specialists) on the plant genus ''Cucurbita'' and closely relate ...
s,
bumble bee
A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only Extant taxon, extant group in the tribe Bombini, though a few extinct r ...
s and other native bees. The
stingless bee
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family A ...
s mentioned by Jefferson are distant relatives of the honey bees, in the genus ''
Melipona
''Melipona'' is a genus of stingless bees, widespread in warm areas of the Neotropics, from Sinaloa and Tamaulipas (México) to Tucumán and Misiones (Argentina). About 70 species are known.Grüter, C. 2020. ''Stingless Bees: Their Behaviour, Ec ...
''.
Nutrition
Honey bees obtain all of their nutritional requirements from a diverse combination of pollen and nectar. Pollen is the only natural protein source for honey bees. Adult worker honey bees consume 3.4–4.3 mg of pollen per day to meet a dry matter requirement of 66–74% protein.
The rearing of one larva requires 125-187.5 mg pollen or 25-37.5 mg protein for proper development.
Dietary proteins are broken down into amino acids, ten of which are considered essential to honey bees: methionine, tryptophan, arginine, lysine, histidine, phenylalanine, isoleucine, threonine, leucine, and valine. Of these amino acids, honey bees require highest concentrations of leucine, isoleucine, and valine, however elevated concentrations of arginine and lysine are required for brood rearing.
In addition to these amino acids, some B vitamins including biotin, folic acid, nicotinamide, riboflavin, thiamine, pantothenate, and most importantly, pyridoxine are required to rear larvae. Pyridoxine is the most prevalent B vitamin found in royal jelly and concentrations vary throughout the foraging season with lowest concentrations found in May and highest concentrations found in July and August. Honey bees lacking dietary pyridoxine were unable to rear brood.
![Bee Collecting Pollen 2004-08-14](https://upload.wikimedia.org/wikipedia/commons/8/81/Bee_Collecting_Pollen_2004-08-14.jpg)
Pollen is also a lipid source for honey bees ranging from 0.8% to 18.9%.
Lipids are metabolized during the brood stage for precursors required for future biosynthesis. Fat-soluble vitamins A, D, E, and K are not considered essential but have shown to significantly improve the number of brood reared.
Honey bees ingest phytosterols from pollen to produce 24-methylenecholesterol and other sterols as they cannot directly synthesize cholesterol from phytosterols. Nurse bees have the ability to selectively transfer sterols to larvae through brood food.
Nectar is collected by foraging worker bees as a source of water and carbohydrates in the form of sucrose. The dominant monosaccharides in honey bee diets are fructose and glucose but the most common circulating sugar in hemolymph is trehalose which is a disaccharide consisting of two glucose molecules. Adult worker honey bees require 4 mg of utilizable sugars per day and larvae require about 59.4 mg of carbohydrates for proper development.
Honey bees require water to maintain osmotic homeostasis, prepare liquid brood food, and to cool the hive through evaporation. A colony's water needs can generally be met by nectar foraging as it has high water content. Occasionally on hot days or when nectar is limited, foragers will collect water from streams or ponds to meet the needs of the hive.
Beekeeping
![Gurian Beekeeper](https://upload.wikimedia.org/wikipedia/commons/2/24/Gurian_Beekeeper.jpg)
The only domesticated species of honey bee are ''A. mellifera'' and ''
A. cerana'', and they are often maintained, fed, and transported by beekeepers. In Japan, where ''A. mellifera'' is vulnerable to local hornets and disease, the Japanese honey bee ''
A. cerana japonica
''Apis cerana japonica'' is a subspecies of the eastern honey bee native to Japan. It is commonly known as the . This subspecies was determined, through an analysis of mitochondrial DNA, to have originally come from the Korean peninsula. They have ...
'' is used in its place. Modern hives also enable beekeepers to transport bees, moving from field to field as the crop needs pollinating and allowing the beekeeper to charge for the pollination services they provide, revising the historical role of the self-employed beekeeper, and favoring large-scale commercial operations. Bees of various types other than honey bees are also domesticated and used for pollination or other means around the world, including ''
Tetragonula iridipennis
The Indian stingless bee or dammar bee, ''Tetragonula iridipennis'', is a species of bee belonging to the family Apidae, subfamily Apinae. It was first described by Frederick Smith in 1854 who found the species in what is now the island of Sri ...
'' in India, the
blue orchard bee
''Osmia lignaria'', commonly known as the orchard mason bee or blue orchard bee, is a megachilid bee that makes nests in reeds and natural holes, creating individual cells for its brood that are separated by mud dividers. Unlike carpenter bees, i ...
for tree nut and fruit pollination in the United States, and a number of species of ''
Bombus
A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only extant group in the tribe Bombini, though a few extinct related gener ...
'' (bumblebees) for pollination in various regions globally, such as
tomato
The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...
es, which are not effectively pollinated by honey bees.
Colony collapse disorder
Primarily in places where western honey bees were imported by humans, periodic collapses in western honey bee populations have occurred at least since the late 19th century. Starting in the first decade of the 21st century, abnormally high die-offs (30–70% of hives) of western honey bee colonies have occurred in North America. This has been dubbed "colony collapse disorder" (CCD) and was at first unexplained. It seems to be caused by a combination of factors rather than a single
pathogen
In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...
or
poison
Poison is a chemical substance that has a detrimental effect to life. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figuratively, with a broa ...
, possibly including
neonicotinoid
Neonicotinoids (sometimes shortened to neonics ) are a class of neuro-active insecticides chemically similar to nicotine, developed by scientists at Shell and Bayer in the 1980s.
The neonicotinoid family includes acetamiprid, clothianidin, dinote ...
pesticides or
Israeli acute paralysis virus
Diseases of the honey bee or abnormal hive conditions include:
Pests and parasites
''Varroa'' mites
''Varroa destructor'' and ''V. jacobsoni'' are parasitic mites that feed on the fat bodies of adult, pupal and larval bees. When the hive ...
.
Parasites
Acarapis woodi
''
Acarapis woodi
''Acarapis woodi'' is an internal parasite affecting honey bees, the symptoms of infestation was originally observed on the Isle of Wight in 1904, but was not species description, described until 1921. ''Acarapis woodi'' mites live and reproduce ...
'' (or "tracheal mites") are parasitic mites which live and reproduce in adult bees' tracheae, or respiratory tubes, piercing the tube walls with their mouthparts to feed on haemolymph. To infest new hosts, the mites must find newly emerged bees; after three days, the bristles (setae) guarding the
spiracles are firm enough to prevent the mites' entry into the tracheae. Mite infestations are known as acarine, and have been called "Isle of Wight disease."
''Galleria mellonella''
Larval stages of the moth ''
G. mellonella'' parasitize both wild and cultivated honey bees, in particular ''Apis mellifera'' and ''Apis cerana''. Eggs are laid within the hive, and the
larva
A larva (; plural larvae ) 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.
The ...
e that hatch tunnel through and destroy the honeycombs that contain bee larva and their honey stores. The tunnels they create are lined with silk, which entangles and starves emerging bees. Destruction of honeycombs also result in honey leaking and being wasted. Both ''G. mellonella'' adults and larvae are possible vectors for pathogens that can infect bees, including the
Israeli acute paralysis virus
Diseases of the honey bee or abnormal hive conditions include:
Pests and parasites
''Varroa'' mites
''Varroa destructor'' and ''V. jacobsoni'' are parasitic mites that feed on the fat bodies of adult, pupal and larval bees. When the hive ...
and the
black queen cell virus
The black queen cell virus (BQCV) is a virus that infects honey bees, specifically ''Apis mellifera'', ''Apis florea'', and ''Apis dorsata''. Infection of the latter two species is more recent and can be attributed to genetic similarity and geogr ...
.
To manage the mite, temperature treatments are possible, but also distorts wax of the honeycombs. Chemical fumigants, particularly CO
2, are also used.
''Varroa'' mites
''Varroa'' mites are arguably the biggest threat to honey bees in the United States. These mites invade hives and reproduce by laying eggs on pupa. The hatching mites eat away at the pupa, causing deformities as well as spreading disease. If not detected and treated early on, the mite population may increase to such an extent that the hive will succumb to the diseases and deformities caused by the mites.
Mite treatment is accomplished by several methods, including treatment strips and acid vaporization.
Bee products
Honey
Honey is the complex substance made when bees ingest nectar, process it, and store the substance into honey combs.
All living species of ''Apis'' have had their honey gathered by indigenous peoples for consumption. ''A. mellifera'' and ''A. cerana'' are the only species that have had their honey harvested for commercial purposes.
Beeswax
Worker bees of a certain age secrete
beeswax
Beeswax (''cera alba'') is a natural wax produced by honey bees of the genus ''Apis''. The wax is formed into scales by eight wax-producing glands in the abdominal segments of worker bees, which discard it in or at the hive. The hive workers ...
from a series of exocrine glands on their abdomens.
They use the wax to form the walls and caps of the comb. As with honey, beeswax is gathered by humans for various purposes such as candle making, waterproofing, soap and cosmetics manufacturing, pharmaceuticals, art, furniture polish and more.
Bee bread
Bees collect
pollen
Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
in their
pollen basket
The pollen basket or corbicula (plural corbiculae) is part of the tibia on the hind legs of certain species of bees. They use the structure in harvesting pollen and carrying it to the nest or hive. Other species of bees have scopae instead.
Ety ...
s and carry it back to the hive.
Worker bees combine pollen, honey and glandular secretions and allow it to ferment in the comb to make bee bread. The fermentation process releases additional nutrients from the pollen and can produce antibiotics and fatty acids which inhibit spoilage. Bee bread is eaten by
nurse bees (younger workers) which produce the protein-rich royal jelly needed by the queen and developing larvae in their hypopharyngeal glands.
In the hive, pollen is used as a
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
source necessary during brood-rearing. In certain environments, excess pollen can be collected from the hives of ''A. mellifera'' and ''A. cerana''. The product is used as a health supplement. It has been used with moderate success as a source of pollen for
hand pollination
Hand pollination, also known as mechanical pollination is a technique that can be used to pollinate plants when natural or open pollination is either undesirable or insufficient.
Method
This method of pollination is done by manually transfer ...
.
Bees as food
Bee brood – the eggs, larvae or
pupa
A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
e of honey bees – is nutritious and seen as a delicacy in countries such as Indonesia,
Mexico, Thailand, and many African countries; it has been consumed since ancient times by the Chinese and Egyptians.
Adult wild honeybees are also consumed as a food in parts of
China
China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
, including
Yunnan
Yunnan , () is a landlocked Provinces of China, province in Southwest China, the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is ...
. According to a worker at a Yunnan-based specialty restaurant, the bees are best served "deep-fried with salt and pepper", and they are "naturally sweet and tasty". Kellie Schmitt of ''
CNN
CNN (Cable News Network) is a multinational cable news channel headquartered in Atlanta, Georgia, U.S. Founded in 1980 by American media proprietor Ted Turner and Reese Schonfeld as a 24-hour cable news channel, and presently owned by the M ...
'' described the dish as one of "
Shanghai
Shanghai (; , , Standard Mandarin pronunciation: ) is one of the four direct-administered municipalities of the People's Republic of China (PRC). The city is located on the southern estuary of the Yangtze River, with the Huangpu River flow ...
's weirdest foods".
Propolis
Propolis is a resinous mixture collected by honey bees from tree buds, sap flows or other botanical sources, which is used as a sealant for unwanted open spaces in the hive. Although propolis is alleged to have health benefits (tincture of Propolis is marketed as a cold and flu remedy), it may cause severe allergic reactions in some individuals. Propolis is also used in wood finishes, and gives a
Stradivarius
A Stradivarius is one of the violins, violas, cellos and other string instruments built by members of the Italian family Stradivari, particularly Antonio Stradivari (Latin: Antonius Stradivarius), during the 17th and 18th centuries. They are co ...
violin its unique red color.
Royal jelly
Royal jelly is a honey bee secretion used to nourish the larvae. It is marketed for its alleged but unsupported claims of health benefits. On the other hand, it may cause severe allergic reactions in some individuals.
Sexes and castes
Honey bees have three
caste
Caste is a form of social stratification characterised by endogamy, hereditary transmission of a style of life which often includes an occupation, ritual status in a hierarchy, and customary social interaction and exclusion based on cultura ...
s:
drones,
workers
The workforce or labour force is a concept referring to the pool of human beings either in employment or in unemployment. It is generally used to describe those working for a single company or industry, but can also apply to a geographic regio ...
, and
queens
Queens is a borough of New York City, coextensive with Queens County, in the U.S. state of New York. Located on Long Island, it is the largest New York City borough by area. It is bordered by the borough of Brooklyn at the western tip of Long ...
.
Drones are male, while workers and queens are female.
[
]
Drones
Drones are typically haploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
, having only one set of chromosome
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s, and primarily exist for the purpose of reproduction
Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual or ...
.[ They are produced by the queen if she chooses not to fertilize an egg or by an unfertilized laying worker. There are rare instances of diploid drone larvae. This phenomenon usually arises when there are more than two generations of brother-sister mating. Sex determination in honey bees is initially due to a single locus, called the complementary sex determiner (''csd'') gene. In developing bees, if the conditions are that the individual is heterozygous for the ''csd'' gene, they will develop into females. If the conditions are so that the individual is ]hemizygous
Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism.
Mo ...
or homozygous for the ''csd'' gene, they will develop into males. The instances where the individual is homozygous at this gene are the instances of diploid males. Drones take 24 days to develop, and may be produced from summer through to autumn, numbering as many as 500 per hive.[ They are expelled from the hive during the winter months when the hive's primary focus is warmth and food conservation.][ Drones have large eyes used to locate queens during mating flights. They do not defend the hive or kill intruders, and do not have a ]stinger
A stinger (or sting) is a sharp organ found in various animals (typically insects and other arthropods) capable of injecting venom, usually by piercing the epidermis of another animal.
An insect sting is complicated by its introduction of v ...
.[
]
Workers
Workers have two sets of chromosomes. They are produced from an egg that the queen has selectively fertilized from stored sperm. Workers typically develop in 21 days. A typical colony may contain as many as 60,000 worker bees.[ Workers exhibit a wider range of behaviors than either queens or drones. Their duties change with age in the following order (beginning with cleaning out their own cell after eating through their capped brood cell): feed brood, receive nectar, clean hive, guard duty, and foraging.] Some workers engage in other specialized behaviors, such as "undertaking" (removing corpses of their nestmates from inside the hive).[
Workers have morphological specializations, including the ]pollen basket
The pollen basket or corbicula (plural corbiculae) is part of the tibia on the hind legs of certain species of bees. They use the structure in harvesting pollen and carrying it to the nest or hive. Other species of bees have scopae instead.
Ety ...
''(corbicula)'', abdominal glands that produce beeswax, brood-feeding glands, and barbs on the sting. Under certain conditions (for example, if the colony becomes queenless), a worker may develop ovaries.
Worker honey bees perform different behavioural tasks that cause them to be exposed to different local environments. The gut microbial composition of workers varies according to the landscape and plant species they forage
Forage is a plant material (mainly plant leaves and stems) eaten by grazing livestock. Historically, the term ''forage'' has meant only plants eaten by the animals directly as pasture, crop residue, or immature cereal crops, but it is also used m ...
, such as differences in rapeseed
Rapeseed (''Brassica napus ''subsp.'' napus''), also known as rape, or oilseed rape, is a bright-yellow flowering member of the family Brassicaceae (mustard or cabbage family), cultivated mainly for its oil-rich seed, which naturally contains a ...
crops,[ and with different hive tasks, such as nursing or food processing.][
]
Queens
Queen honey bees are created when worker bees feed a single female larva an exclusive diet of a food called "royal jelly
Royal jelly is a honey bee secretion that is used in the nutrition of larvae and adult queens. It is secreted from the glands in the hypopharynx of nurse bees, and fed to all larvae in the colony, regardless of sex or caste.Graham, J. (ed.) (19 ...
".[ Queens are produced in oversized cells and develop in only 16 days; they differ in physiology, morphology, and behavior from worker bees. In addition to the greater size of the queen, she has a functional set of ovaries, and a spermatheca, which stores and maintains sperm after she has mated. ''Apis'' queens practice ]polyandry
Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives" ...
, with one female mating with multiple males. The highest documented mating frequency for an ''Apis'' queen is in ''Apis nigrocincta
''Apis nigrocincta'' is a species of honey bee that inhabits the Philippine island of Mindanao as well as the Indonesian islands of Sangihe and Sulawesi. The species is known to have queens with the highest mating frequencies of any species of ...
'', where queens mate with an extremely high number of males with observed numbers of different matings ranging from 42 to 69 drones per queen. The sting of queens is not barbed like a worker's sting, and queens lack the glands that produce beeswax. Once mated, queens may lay up to 2,000 eggs per day.[ They produce a variety of pheromones that regulate behavior of workers, and helps swarms track the queen's location during the swarming.][
]
Queen-worker conflict
When a fertile female worker produces drones, a conflict arises between her interests and those of the queen. The worker shares half her genes with the drone and one-quarter with her brothers, favouring her offspring over those of the queen. The queen shares half her genes with her sons and one-quarter with the sons of fertile female workers. This pits the worker against the queen and other workers, who try to maximize their reproductive fitness
Fitness (often denoted w or ω in population genetics models) is the quantitative representation of individual reproductive success. It is also equal to the average contribution to the gene pool of the next generation, made by the same individua ...
by rearing the offspring most related to them. This relationship leads to a phenomenon known as "worker policing". In these rare situations, other worker bees in the hive who are genetically more related to the queen's sons than those of the fertile workers will patrol the hive and remove worker-laid eggs. Another form of worker-based policing is aggression toward fertile females. Some studies have suggested a queen pheromone which may help workers distinguish worker- and queen-laid eggs, but others indicate egg viability as the key factor in eliciting the behavior. Worker policing is an example of forced altruism, where the benefits of worker reproduction are minimized and that of rearing the queen's offspring maximized.
In very rare instances workers subvert the policing mechanisms of the hive, laying eggs which are removed at a lower rate by other workers; this is known as anarchic syndrome. Anarchic workers can activate their ovaries at a higher rate and contribute a greater proportion of males to the hive. Although an increase in the number of drones would decrease the overall productivity of the hive, the reproductive fitness of the drones' mother would increase. Anarchic syndrome is an example of selection working in opposite directions at the individual and group levels for the stability of the hive.
Under ordinary circumstances the death (or removal) of a queen increases reproduction in workers, and a significant proportion of workers will have active ovaries in the absence of a queen. The workers of the hive produce a last batch of drones before the hive eventually collapses. Although during this period worker policing is usually absent, in certain groups of bees it continues.
According to the strategy of kin selection
Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin altruism can look like Altruism in animals, altruistic behavio ...
, worker policing is not favored if a queen does not mate multiple times. Workers would be related by three-quarters of their genes, and the difference in relationship between sons of the queen and those of the other workers would decrease. The benefit of policing is negated, and policing is less favored. Experiments confirming this hypothesis have shown a correlation between higher mating rates and increased rates of worker policing in many species of social hymenoptera
Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
.
Defense
All honey bees live in colonies where the workers sting
Sting may refer to:
* Stinger or sting, a structure of an animal to inject venom, or the injury produced by a stinger
* Irritating hairs or prickles of a stinging plant, or the plant itself
Fictional characters and entities
* Sting (Middle-eart ...
intruders as a form of defense, and alarmed bees release a 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 like hormones outside the body of the secreting individual, to affect the behavio ...
that stimulates the attack response in other bees. The different species of honey bees are distinguished from all other bee species by the possession of small barbs on the sting, but these barbs are found only in the worker bees.
The sting apparatus, including the barbs, may have evolved specifically in response to predation by vertebrates, as the barbs do not usually function (and the sting apparatus does not detach) unless the sting is embedded in fleshy tissue. While the sting can also penetrate the membranes between joints in the exoskeleton of other insects (and is used in fights between queens), in the case of ''Apis cerana japonica'', defense against larger insects such as predatory wasps (e.g. Asian giant hornet
The Asian giant hornet (''Vespa mandarinia'') or northern giant hornet, including the color form referred to as the Japanese giant hornet, is the world's largest hornet. It is native to temperate and tropical East Asia, South Asia, Mainland Sout ...
) is usually performed by surrounding the intruder with a mass of defending worker bees, which vibrate their muscles vigorously to raise the temperature of the intruder to a lethal level ("balling"). Previously, heat alone was thought to be responsible for killing intruding wasps, but recent experiments have demonstrated the increased temperature in combination with increased carbon dioxide levels within the ball produce the lethal effect. This phenomenon is also used to kill a queen perceived as intruding or defective, an action known to beekeepers as 'balling the queen', named for the ball of bees formed.
Defense can vary based on the habitat of the bee. In the case of those honey bee species with open combs (e.g., ''A. dorsata''), would-be predators are given a warning signal that takes the form of a "wave
In physics, mathematics, and related fields, a wave is a propagating dynamic disturbance (change from equilibrium) of one or more quantities. Waves can be periodic, in which case those quantities oscillate repeatedly about an equilibrium (res ...
" that spreads as a ripple across a layer of bees densely packed on the surface of the comb when a threat is perceived, and consists of bees momentarily arching their bodies and flicking their wings. In cavity dwelling species such as ''Apis cerana
''Apis cerana'', the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South, Southeast and East Asia. This species is the sister species of ''Apis koschevnikovi'' and both are in the same subgenus as th ...
'', ''Apis mellifera
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
'', and ''Apis nigrocincta
''Apis nigrocincta'' is a species of honey bee that inhabits the Philippine island of Mindanao as well as the Indonesian islands of Sangihe and Sulawesi. The species is known to have queens with the highest mating frequencies of any species of ...
'', entrances to these cavities are guarded and checked for intruders in incoming traffic. Another act of defense against nest invaders, particularly wasps, is "body shaking," a violent and pendulum like swaying of the abdomen, performed by worker bees.
A 2020 study of ''Apis cerana
''Apis cerana'', the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South, Southeast and East Asia. This species is the sister species of ''Apis koschevnikovi'' and both are in the same subgenus as th ...
'' in Vietnam
Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
found that they use feces
Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
and even human urine to defend their hives against raids by hornets (''Vespa soror
Vespa () is an Italian luxury brand of scooters and mopeds manufactured by Piaggio. The name means wasp in Italian. The Vespa has evolved from a single model motor scooter manufactured in 1946 by Piaggio & Co. S.p.A. of Pontedera, Italy to ...
''), a strategy not replicated by their European and North American counterparts, though collection and use of feces in nest construction is well-known in stingless bee
Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family A ...
s.
Venom
The stings of honey bees are barbed and therefore embed themselves into the sting site, and the sting apparatus has its own musculature and ganglion which keep delivering venom even after detachment. The gland which produces the alarm pheromone is also associated with the sting apparatus. The embedded stinger continues to emit additional alarm pheromone after it has torn loose; other defensive workers are thereby attracted to the sting site. The worker dies after the sting becomes lodged and is subsequently torn loose from the bee's abdomen. The honey bee's venom, known as apitoxin
Apitoxin or bee venom is the venom produced by the honey bee. It is a cytotoxic and hemotoxic bitter colorless liquid containing proteins, which may produce local inflammation. It may have similarities to sea nettle toxin.
Components
Bee venom i ...
, carries several active components, the most abundant of which is melittin
Melittin is the main component (40–60% of the dry weight) and the major pain producing substance of honeybee (''Apis mellifera'') venom. Melittin is a basic peptide consisting of 26 amino acids.
Function
The principal function of melittin a ...
, and the most biologically active are enzyme
Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. A ...
s, particularly phospholipase A2
The enzyme phospholipase A2 (EC 3.1.1.4, PLA2, systematic name phosphatidylcholine 2-acylhydrolase) catalyse the cleavage of fatty acids in position 2 of phospholipids, hydrolyzing the bond between the second fatty acid “tail” and the glyce ...
.
Honey bee venom is under laboratory and clinical research
Clinical research is a branch of healthcare science that determines the safety and effectiveness ( efficacy) of medications, devices, diagnostic products and treatment regimens intended for human use. These may be used for prevention, treatm ...
for its potential properties and uses in reducing risks for adverse event
An adverse event (AE) is any untoward medical occurrence in a patient or clinical investigation subject administered a pharmaceutical product and which does not necessarily have a causal relationship with this treatment. An adverse event can ther ...
s from bee venom therapy
A therapy or medical treatment (often abbreviated tx, Tx, or Tx) is the attempted remediation of a health problem, usually following a medical diagnosis.
As a rule, each therapy has indications and contraindications. There are many different ...
, rheumatoid arthritis
Rheumatoid arthritis (RA) is a long-term autoimmune disorder that primarily affects joints. It typically results in warm, swollen, and painful joints. Pain and stiffness often worsen following rest. Most commonly, the wrist and hands are involv ...
, and use as an immunotherapy
Immunotherapy or biological therapy is the treatment of disease by activating or suppressing the immune system. Immunotherapies designed to elicit or amplify an immune response are classified as ''activation immunotherapies,'' while immunotherap ...
for protection against allergies
Allergies, also known as allergic diseases, refer a number of conditions caused by the hypersensitivity of the immune system to typically harmless substances in the environment. These diseases include hay fever, food allergies, atopic derma ...
from insect stings. Bee venom products are marketed in many countries, but, as of 2018, there are no approved clinical uses for these products which carry various warnings for potential allergic reactions.
Competition
With an increased number of honey bees in a specific area due to beekeeping, Western honey bees (as an invasive species
An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
) and native wild bees often have to compete for the limited habitat and food sources available, and Western honey bees may become defensive in response to the seasonal arrival of competition from other colonies, particularly Africanized bees which may be on the offence and defence year round due to their tropical origin.
Communication
Honey bees are known to communicate through many different chemicals and odors, as is common in insects. They also rely on a sophisticated dance language that conveys information about the distance and direction to a specific location (typically a nutritional source, e.g., flowers or water). The dance language is also used during the process of reproductive fission, or swarming, when scouts communicate the location and quality of nesting sites.
The details of the signalling being used vary from species to species; for example, the two smallest species, ''Apis andreniformis
''Apis andreniformis'', or the black dwarf honey bee, is a relatively rare species of honey bee whose native habitat is the tropical and subtropical regions of Southeast Asia.
''A. andreniformis'' was the fifth honey bee species to be described ...
'' and ''A. florea'', dance on the upper surface of the comb, which is horizontal (not vertical, as in other species), and worker bees orient the dance in the actual compass direction of the resource to which they are recruiting.
Carniolan honey bee
The Carniolan honey bee (''Apis mellifera carnica'', Pollmann) is a subspecies of the western honey bee. The Carniolan honey bee is native to Slovenia, southern Austria, and parts of Croatia, Bosnia and Herzegovina, Montenegro, Serbia, Hungary, ...
s (''Apis mellifera carnica'') use their antennae asymmetrically for social interactions, with a strong lateral preference to use their right antennae.
There has been speculation as to honey bee consciousness
Consciousness, at its simplest, is sentience and awareness of internal and external existence. However, the lack of definitions has led to millennia of analyses, explanations and debates by philosophers, theologians, linguisticians, and scien ...
. While honey bees lack the parts of the brain that a human being uses for consciousness like the cerebral cortex or even the cerebrum itself, when those parts of a human brain are damaged, the midbrain seems able to provide a small amount of consciousness. Honey bees have a tiny structure that appears similar to a human midbrain, so if it functions the same way they may possibly be able to achieve a small amount of simple awareness of their bodies.
Symbolism
The bee was used as a symbol of government by Emperor Napoleon I
Napoleon Bonaparte ; it, Napoleone Bonaparte, ; co, Napulione Buonaparte. (born Napoleone Buonaparte; 15 August 1769 – 5 May 1821), later known by his regnal name Napoleon I, was a French military commander and political leader who ...
of France. Both the Hindu
Hindus (; ) are people who religiously adhere to Hinduism.Jeffery D. Long (2007), A Vision for Hinduism, IB Tauris, , pages 35–37 Historically, the term has also been used as a geographical, cultural, and later religious identifier for ...
''Atharva Veda
The Atharva Veda (, ' from ' and ''veda'', meaning "knowledge") is the "knowledge storehouse of ''atharvāṇas'', the procedures for everyday life".Laurie Patton (2004), Veda and Upanishad, in ''The Hindu World'' (Editors: Sushil Mittal and G ...
'' and the ancient Greeks associated lips anointed with honey with the gift of eloquence and even of prescience. The priestess at Delphi
Delphi (; ), in legend previously called Pytho (Πυθώ), in ancient times was a sacred precinct that served as the seat of Pythia, the major oracle who was consulted about important decisions throughout the ancient classical world. The oracle ...
was the "Delphic Bee".
The ''Quran
The Quran (, ; Standard Arabic: , Classical Arabic, Quranic Arabic: , , 'the recitation'), also romanized Qur'an or Koran, is the central religious text of Islam, believed by Muslims to be a revelation in Islam, revelation from God in Islam, ...
'' has a Sura
A ''surah'' (; ar, سورة, sūrah, , ), is the equivalent of "chapter" in the Qur'an. There are 114 ''surahs'' in the Quran, each divided into '' ayats'' (verses). The chapters or ''surahs'' are of unequal length; the shortest surah ('' Al-K ...
(chapter) titled " The Bee". It is named after honey bees, and contains a comparison of the industry and adaptability of honey bees to the industry of man.
In ancient Egyptian mythology, honey bees were believed to be born from the tears of the Sun God
A solar deity or sun deity is a deity who represents the Sun, or an aspect of it. Such deities are usually associated with power and strength. Solar deities and Sun worship can be found throughout most of recorded history in various forms. The ...
, Ra.
A community of honey bees has often been employed by political theorists as a model of human society, from Aristotle and Plato to Virgil. Honey bees, signifying immortality and resurrection, were royal emblems of the Merovingians
The Merovingian dynasty () was the ruling family of the Franks from the middle of the 5th century until 751. They first appear as "Kings of the Franks" in the Roman army of northern Gaul. By 509 they had united all the Franks and northern Gauli ...
. The state of Utah
Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
is called the "Beehive State", the state emblem is the beehive, the state insect is the honey bee, and a beehive and the word "industry" appear on both the state flag and seal.
Gallery
File:Bienenkoenigin.JPG, A coloured dot applied by a beekeeper identifies a queen
Queen or QUEEN may refer to:
Monarchy
* Queen regnant, a female monarch of a Kingdom
** List of queens regnant
* Queen consort, the wife of a reigning king
* Queen dowager, the widow of a king
* Queen mother, a queen dowager who is the mother ...
Western honey bee
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
File:Honeybee02.jpg, Western honey bee
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
foragers loaded with pollen on the hive landing board
File:Cerana.jpg, Eastern honey bee
''Apis cerana'', the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South Asia, South, Southeast Asia, Southeast and East Asia. This species is the sister taxon, sister species of ''Apis koschevnikovi ...
(''A. cerana'') in Hong Kong
Hong Kong ( (US) or (UK); , ), officially the Hong Kong Special Administrative Region of the People's Republic of China ( abbr. Hong Kong SAR or HKSAR), is a city and special administrative region of China on the eastern Pearl River Delt ...
File:ApisDorsataHive.jpg, Giant honey bee
''Apis dorsata'', the giant honey bee, सिङ्गुस in Nepali, is a honey bee of South and Southeast Asia, found mainly in forested areas such as the Terai of Nepal. They are typically around long. Nests are mainly built in exposed pla ...
(''A. dorsata'')
File:Pollinationn.jpg, Western honey bee
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
visiting flowers
File:Natural Beehive and Honeycombs.jpg, A colony of giant honey bees (''A. dorsata'') on their comb
File:Bee-apis.jpg, Western honey bee
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
File:Honey bee on rock rose.jpg, Western honey bee
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
on rock rose (''Cistus
''Cistus'' (from the Greek ''kistos'') is a genus of flowering plants in the rockrose family Cistaceae, containing about 20 species (Ellul ''et al.'' 2002). They are perennial shrubs found on dry or rocky soils throughout the Mediterranean region ...
'') in Oakland, California
File:Honeybee pollen turnips.jpg, Western honey bee
The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying", ...
collecting pollen from turnip blossoms in Eastern Oklahoma
In the U.S. state of Oklahoma, Eastern Oklahoma is an amorphous area roughly defined as east of Oklahoma City and/or east of I-35. The Oklahoma Department of Tourism and Recreation established regional designations for the various parts of the st ...
File:Green Bee-eater Merops orientalis feeding on Honey-bee by Dr. Raju Kasambe DSCN1640 (1).jpg, A predator, the green bee-eater
The Asian green bee-eater (''Merops orientalis''), also known as little green bee-eater, and green bee-eater in Sri Lanka, is a near passerine bird in the bee-eater family. It is resident but prone to seasonal movements and is found widely distri ...
File:A natural beehive in Chandigarh.jpg, A wild colony in Chandigarh
See also
* Bees and toxic chemicals
Bees can suffer serious effects from toxic chemicals in their environments. These include various synthetic chemicals, particularly insecticides, as well as a variety of naturally occurring chemicals from plants, such as ethanol resulting from ...
* Honey bee life cycle
The honey bee life cycle, here referring exclusively to the domesticated Western honey bee, depends greatly on their social structure.
Honey bee colony life
Unlike a bumble bee colony or a paper wasp colony, the life of a honey bee colony is pe ...
* Honey bee starvation
Honey bee starvation is a problem for bees and beekeepers. Starvation may be caused by unfavorable weather, disease, long distance transportation or depleting food reserve. Over-harvesting of honey (and the lack of supplemental feeding) is the fore ...
* ''More than Honey
More or Mores may refer to:
Computing
* MORE (application), outline software for Mac OS
* more (command), a shell command
* MORE protocol, a routing protocol
* Missouri Research and Education Network
Music Albums
* ''More!'' (album), by Booka ...
'' – a 2012 Swiss documentary film about honey bees
*Melittology
Melittology (from Greek , ''melitta'', "bee"; and ''-logia'') is a branch of entomology concerning the scientific study of bees. It may also be called apicology. Melittology covers the species found in the clade Anthophila within the superfamil ...
, the study of bees
Notes
References
External links
*
Could a Mushroom Save a Honeybee?
' Documentary produced by Oregon Field Guide
''Oregon Field Guide'' is a weekly television program produced by Oregon Public Broadcasting focusing on recreation, the outdoors, and environmental issues in the state of Oregon. The show has become part of the Oregon zeitgeist. Steve Amen is th ...
, 14 October 2015
{{DEFAULTSORT:Honey Bee
Apis (genus)
Articles containing video clips
Beekeeping
Bees
Extant Oligocene first appearances
Insects in culture
Bees described in 1758
Taxa named by Carl Linnaeus