Bees are winged
insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s closely related to
wasp
A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s and
ant
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s, known for their roles in
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 ...
and, in the case of the best-known bee species, 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", ...
, for producing
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 ...
. Bees are a
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
lineage within the superfamily
Apoidea
The superfamily (zoology), superfamily Apoidea is a major group within the Hymenoptera, which includes two traditionally recognized lineages, the "sphecidae, sphecoid" wasps, and the bees. Molecular phylogeny demonstrates that the bees arose from ...
. They are presently considered a
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, ...
, called Anthophila. There are over 16,000 known species of bees in seven recognized
biological families.
[ Some speciesincluding ]honey bee
A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current co ...
s, 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 ...
slive socially
Social organisms, including human(s), live collectively in interacting populations. This interaction is considered social whether they are aware of it or not, and whether the exchange is voluntary or not.
Etymology
The word "social" derives from ...
in colonies while most species (>90%)including mason bee
Mason bee is a name now commonly used for species of bees in the genus ''Osmia'', of the family Megachilidae. Mason bees are named for their habit of using mud or other "masonry" products in constructing their nests, which are made in naturally ...
s, carpenter bee
Carpenter bees are species in the genus ''Xylocopa'' of the subfamily Xylocopinae. The genus includes some 500 bees in 31 subgenera. The common name "carpenter bee" derives from their nesting behavior; nearly all species burrow into hard plant m ...
s, leafcutter bees
The genus ''Megachile'' is a cosmopolitan group of solitary bees, often called leafcutter bees or leafcutting bees; it also includes the called resin bees and mortar bees. While other genera within the family Megachilidae may chew leaves or pet ...
, and sweat beesare solitary.
Bees are found on every continent except Antarctica
Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...
, in every habitat on the planet that contains insect-pollinated flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s. The most common bees in the Northern Hemisphere
The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
are the Halictidae
Halictidae is the second-largest family of bees (clade Anthophila) with nearly 4,500 species. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every contine ...
, or sweat bees, but they are small and often mistaken for wasps or flies. Bees range in size from tiny 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 ...
species, whose workers are less than long, to ''Megachile pluto
''Megachile pluto'', also known as Wallace's giant bee or (), is a large resin bee found in Indonesia. With a wingspan of it is the largest known living bee species. It was believed to be extinct until several specimens were discovered in 1981 ...
'', the largest species of leafcutter bee, whose females can attain a length of .
Bees feed on nectar
Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
and 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 ...
, the former primarily as an energy source and the latter primarily for 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 ...
and other nutrients. Most pollen is used as food for their 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. Vertebrate predators of bees include primates
Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
and birds such as bee-eater
The bee-eaters are a group of non-passerine birds in the family Meropidae, containing three genera and thirty species. Most species are found in Africa and Asia, with a few in southern Europe, Australia, and New Guinea. They are characterised by ...
s; insect predators include beewolves and dragonflies
A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
.
Bee 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 ...
is important both ecologically and commercially
Commerce is the large-scale organized system of activities, functions, procedures and institutions directly and indirectly related to the exchange (buying and selling) of goods and services among two or more parties within local, regional, nation ...
, and the decline in wild bees has increased the value of pollination by commercially managed hives of honey bees. The analysis of 353 wild bee and hoverfly species across Britain from 1980 to 2013 found the insects have been lost from a quarter of the places they inhabited in 1980.
Human beekeeping
Beekeeping (or apiculture) is the maintenance of bee colonies, commonly in man-made beehives. Honey bees in the genus '' Apis'' are the most-commonly-kept species but other honey-producing bees such as ''Melipona'' stingless bees are also kept. ...
or apiculture (meliponiculture for stingless bees) has been practised for millennia, since at least the times of Ancient Egypt and Ancient Greece
Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity ( AD 600), that comprised a loose collection of cult ...
. Bees have appeared in mythology and folklore, through all phases of art and literature from ancient times to the present day, although primarily focused in the Northern Hemisphere
The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
where beekeeping
Beekeeping (or apiculture) is the maintenance of bee colonies, commonly in man-made beehives. Honey bees in the genus '' Apis'' are the most-commonly-kept species but other honey-producing bees such as ''Melipona'' stingless bees are also kept. ...
is far more common. In Mesoamerica
Mesoamerica is a historical region and cultural area in southern North America and most of Central America. It extends from approximately central Mexico through Belize, Guatemala, El Salvador, Honduras, Nicaragua, and northern Costa Rica. W ...
, the Mayans
The Maya peoples () are an ethnolinguistic group of indigenous peoples of Mesoamerica. The ancient Maya civilization was formed by members of this group, and today's Maya are generally descended from people who lived within that historical reg ...
have practiced large-scale intensive meliponiculture since pre-Columbian times.
Evolution
The immediate ancestors of bees were stinging wasps in the family Crabronidae
The Crabronidae are a large paraphyletic group (nominally a family) of wasps, including nearly all of the species formerly comprising the now-defunct superfamily Sphecoidea. It collectively includes well over 200 genera, containing well over 9 ...
, which were predators
Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
of other insects. The switch from insect prey to pollen may have resulted from the consumption of prey insects which were flower visitors and were partially covered with pollen when they were fed to the wasp larvae. This same 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 ...
ary scenario may have occurred within the vespoid wasps, where the pollen wasps evolved from predatory ancestors. The oldest non-compression bee fossil is found in New Jersey amber
New Jersey Amber, sometimes called Raritan amber, is amber found in the Raritan and Magothy Formations of the Central Atlantic (Eastern) coast of the United States. It is dated to the Late Cretaceous, Turonian age, based on pollen analysis of th ...
, '' Cretotrigona prisca'', a corbiculate bee of Cretaceous age (~65 mya). A fossil from the early Cretaceous
The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma.
Geology
Pro ...
(~100 mya), '' Melittosphex burmensis'', was initially considered "an extinct lineage of pollen-collecting Apoidea sister
A sister is a woman or a girl who shares one or more parents with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to refer to ...
to the modern bees", but subsequent research has rejected the claim that ''Melittosphex'' is a bee, or even a member of the superfamily Apoidea
The superfamily (zoology), superfamily Apoidea is a major group within the Hymenoptera, which includes two traditionally recognized lineages, the "sphecidae, sphecoid" wasps, and the bees. Molecular phylogeny demonstrates that the bees arose from ...
to which bees belong, instead treating the lineage as ''incertae sedis
' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'' within the Aculeata
Aculeata is a subclade of Hymenoptera containing ants, bees, and stinging wasps. The name is a reference to the defining feature of the group, which is the modification of the ovipositor into a stinger. However, many members of the group cannot ...
. By 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' ...
(~45 mya) there was already considerable diversity among eusocial bee lineages.
The highly 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 ...
corbiculate Apidae appeared roughly 87 Mya, and the Allodapini (within the Apidae) around 53 Mya.
The Colletidae
The Colletidae are a family of bees, and are often referred to collectively as plasterer bees or polyester bees, due to the method of smoothing the walls of their nest cells with secretions applied with their mouthparts; these secretions dry int ...
appear as fossils only from the late 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 ...
(~25 Mya) to early Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
.
The Melittidae
Melittidae is a small bee family, with over 200 described species in three subfamilies. The family has a limited distribution, with all described species restricted to Africa and the northern temperate zone.
Fossil melittids have been found oc ...
are known from ''Palaeomacropis eocenicus'' in the Early Eocene
In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresian i ...
.
The Megachilidae
Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a ''scopa'') is restricted to the ventral surface of the abdomen (rather than mostly or exclusively on the hind legs as in other b ...
are known from trace fossils (characteristic leaf cuttings) from the Middle Eocene
The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "dawn ...
.
The Andrenidae
The Andrenidae (commonly known as mining bees) are a large, nearly cosmopolitan family of solitary, ground-nesting bees. Most of the family's diversity is located in temperate or arid areas (warm temperate xeric). It includes some enormous gene ...
are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale.
The Halictidae
Halictidae is the second-largest family of bees (clade Anthophila) with nearly 4,500 species. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every contine ...
first appear in the Early Eocene with species found in amber. The Stenotritidae
The Stenotritidae is the smallest of all formally recognised bee families, with only 21 species in two genera, all of them restricted to Australia. Historically, they were generally considered to belong in the family Colletidae, but the stenotrit ...
are known from fossil brood cells of Pleistocene
The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
age.
Coevolution
The earliest animal-pollinated flowers were shallow, cup-shaped blooms pollinated
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, ...
by insects such as beetle
Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s, so the syndrome of insect pollination was well established before the first appearance of bees. The novelty is that bees are specialized as pollination agents, with behavioral and physical modifications that specifically enhance pollination, and are the most efficient pollinating insects. In a process of coevolution
In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
, flowers developed floral rewards such as nectar
Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
and longer tubes, and bees developed longer tongues to extract the nectar.[ Bees also developed structures known as scopal hairs and ]pollen baskets
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.
Et ...
to collect and carry pollen. The location and type differ among and between groups of bees. Most species have scopal hairs on their hind legs or on the underside of their abdomens. Some species in the family Apidae have pollen baskets
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.
Et ...
on their hind legs, while very few lack these and instead collect pollen in their crops. The appearance of these structures drove the adaptive radiation
In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
of the angiosperms
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
, and, in turn, bees themselves. Bees coevolved
In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
not only with flowers but it is believed that some species coevolved with mites. Some provide tufts of hairs called acarinaria that appear to provide lodgings for mites; in return, it is believed that mites eat fungi that attack pollen, so the relationship in this case may be mutualistic.
Phylogeny
External
This phylogenetic tree
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 based on Debevic ''et al'', 2012, which used molecular phylogeny to demonstrate that the bees (Anthophila
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 monophyletic lineage within the superfami ...
) arose from deep within the Crabronidae
The Crabronidae are a large paraphyletic group (nominally a family) of wasps, including nearly all of the species formerly comprising the now-defunct superfamily Sphecoidea. It collectively includes well over 200 genera, containing well over 9 ...
, which is therefore 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 ...
. The placement of the Heterogynaidae is uncertain. The small subfamily Mellininae
Mellininae is a very small subfamily of wasps, comprising only 17 described species in two genera. Occasionally this group has been treated as a separate family (Mellinidae), but has been more typically considered a subfamily within the all-inclu ...
was not included in this analysis.
Internal
This cladogram of the bee families is based on Hedtke et al., 2013, which places the former families Dasypodaidae and Meganomiidae as subfamilies inside the Melittidae. English names, where available, are given in parentheses.
Characteristics
Bees differ from closely related groups such as wasps by having branched or plume-like seta
In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms.
Animal setae
Protostomes
Annelid setae are stiff bristles present on the body. Th ...
e (hairs), combs on the forelimbs for cleaning their antennae, small anatomical differences in limb structure, and the venation of the hind wings; and in females, by having the seventh dorsal abdominal plate divided into two half-plates.
Bees have the following characteristics:
* A pair of large compound eyes
A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which distin ...
which cover much of the surface of the head. Between and above these are three small simple eyes (ocelli
A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
) which provide information on light intensity.
* The antennae usually have 13 segments in males and 12 in females, and are geniculate, having an elbow joint part way along. They house large numbers of sense organs that can detect touch (mechanoreceptors), smell and taste; and small, hairlike mechanoreceptors that can detect air movement so as to "hear" sounds.
* The mouthparts are adapted for both chewing and sucking by having both a pair of mandibles
In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
and a long proboscis
A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
for sucking up nectar.
* The thorax has three segments, each with a pair of robust legs, and a pair of membranous wings on the hind two segments. The front legs of corbiculate bees bear combs for cleaning the antennae, and in many species the hind legs bear pollen baskets, flattened sections with incurving hairs to secure the collected pollen. The wings are synchronised in flight, and the somewhat smaller hind wings connect to the forewings by a row of hooks along their margin which connect to a groove in the forewing.
* The abdomen has nine segments, the hindermost three being modified into the sting.[
]
The largest species of bee is thought to be Wallace's giant bee ''Megachile pluto
''Megachile pluto'', also known as Wallace's giant bee or (), is a large resin bee found in Indonesia. With a wingspan of it is the largest known living bee species. It was believed to be extinct until several specimens were discovered in 1981 ...
'', whose females can attain a length of . The smallest species may be dwarf stingless bees in the tribe Meliponini
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 ...
whose workers are less than in length.
Sociality
Haplodiploid breeding system
According to inclusive fitness
In evolutionary biology, inclusive fitness is one of two metrics of evolutionary success as defined by W. D. Hamilton in 1964:
* Personal fitness is the number of offspring that an individual begets (regardless of who rescues/rears/supports them ...
theory, organisms can gain fitness not just through increasing their own reproductive output, but also that of close relatives. In evolutionary terms, individuals should help relatives when ''Cost < Relatedness * Benefit''. The requirements for eusociality are more easily fulfilled by haplodiploid
Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky.
Haplodiploidy determines the se ...
species such as bees because of their unusual relatedness structure.
In haplodiploid species, females develop from fertilized eggs and males from unfertilized eggs. Because a male is 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 ...
(has only one copy of each gene), his daughters (which 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 ...
, with two copies of each gene) share 100% of his genes and 50% of their mother's. Therefore, they share 75% of their genes with each other. This mechanism of sex determination gives rise to what W. D. Hamilton
William Donald Hamilton (1 August 1936 – 7 March 2000) was a British evolutionary biologist, recognised as one of the most significant evolutionary theorists of the 20th century.
Hamilton became known for his theoretical work expounding a ...
termed "supersisters", more closely related to their sisters than they would be to their own offspring. Workers often do not reproduce, but they can pass on more of their genes by helping to raise their sisters (as queens) than they would by having their own offspring (each of which would only have 50% of their genes), assuming they would produce similar numbers. This unusual situation has been proposed as an explanation of the multiple (at least 9) evolutions of eusociality within 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 ...
.
Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such as termites
Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
are not haplodiploid. Conversely, all bees are haplodiploid but not all are eusocial, and among eusocial species many queens mate with multiple males, creating half-sisters that share only 25% of each-other's genes. But, monogamy (queens mating singly) is the ancestral state for all eusocial species so far investigated, so it is likely that haplodiploidy contributed to the evolution of eusociality in bees.
Eusociality
Bees may be solitary or may live in various types of communities. Eusociality
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 genera ...
appears to have originated from at least three independent origins in halictid bees. The most advanced of these are species with 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 ...
colonies; these are characterised by cooperative brood care and a division of labour
The division of labour is the separation of the tasks in any economic system or organisation so that participants may specialise (specialisation). Individuals, organizations, and nations are endowed with, or acquire specialised capabilities, and ...
into reproductive and non-reproductive adults, plus overlapping generations. This division of labour creates specialized groups within eusocial societies which are called castes
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 ...
. In some species, groups of cohabiting females may be sisters, and if there is a division of labour within the group, they are considered semisocial
Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies.
Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...
. The group is called eusocial if, in addition, the group consists of a mother (the 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 ...
) and her daughters (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 ...
). When the castes are purely behavioural alternatives, with no morphological differentiation other than size, the system is considered primitively eusocial, as in many paper wasp
Paper wasps are Vespidae, vespid wasps that gather fibers from dead wood and plant stems, which they mix with saliva, and use to construct nests made of gray or brown papery material. Some types of paper wasps are also sometimes called umbrell ...
s; when the castes are morphologically discrete, the system is considered highly eusocial.
True honey bees (genus '' Apis'', of which eight species are currently recognized) are highly eusocial, and are among the best known insects. Their colonies are established by swarms, consisting of a queen and several thousand workers. There are 29 subspecies of one of these species, ''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", ...
'', native to Europe, the Middle East, and Africa. Africanized bee
The Africanized bee, also known as the Africanized honey bee and known colloquially as the "killer bee", is a hybrid of the western honey bee (''Apis mellifera''), produced originally by crossbreeding of the East African lowland honey bee (''A. ...
s are a hybrid strain of ''A. mellifera'' that escaped from experiments involving crossing European and African subspecies; they are extremely defensive.
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 highly 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 ...
. They practise mass provisioning
Mass provisioning is a form of parental investment in which an adult insect, most commonly a hymenopteran such as a bee or wasp, stocks all the food for each of her offspring in a small chamber (a "cell") before she lays the egg. This behavior i ...
, with complex nest architecture and perennial colonies also established via swarming.
Many bumblebees are eusocial, similar to the eusocial Vespidae
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 ...
such as hornet
Hornets (insects in the genus ''Vespa'') are the largest of the eusocial wasps, and are similar in appearance to their close relatives yellowjackets. Some species can reach up to in length. They are distinguished from other vespine wasps by th ...
s in that the queen initiates a nest on her own rather than by swarming. Bumblebee colonies typically have from 50 to 200 bees at peak population, which occurs in mid to late summer. Nest architecture is simple, limited by the size of the pre-existing nest cavity, and colonies rarely last more than a year. In 2011, the International Union for Conservation of Nature
The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
set up the Bumblebee Specialist Group to review the threat status of all bumblebee species worldwide using the IUCN Red List
The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
criteria.
There are many more species of primitively eusocial than highly eusocial bees, but they have been studied less often. Most are in the family Halictidae
Halictidae is the second-largest family of bees (clade Anthophila) with nearly 4,500 species. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every contine ...
, or "sweat bees". Colonies are typically small, with a dozen or fewer workers, on average. Queens and workers differ only in size, if at all. Most species have a single season colony cycle, even in the tropics, and only mated females hibernate. A few species have long active seasons and attain colony sizes in the hundreds, such as ''Halictus hesperus
The genus ''Halictus'' is a large assemblage of bee species in the family Halictidae. The genus is divided into 15 subgenera, some of dubious monophyly, containing over 200 species, primarily in the Northern Hemisphere (a few species occur in So ...
''. Some species are eusocial in parts of their range and solitary in others, or have a mix of eusocial and solitary nests in the same population. The orchid bee
The tribe Euglossini, in the subfamily Apinae, commonly known as orchid bees or euglossine bees, are the only group of corbiculate bees whose non-parasitic members do not all possess eusocial behavior.
Description
Most of the tribe's species ...
s (Apidae) include some primitively eusocial species with similar biology. Some allodapine bees (Apidae) form primitively eusocial colonies, with progressive provisioning Progressive provisioning is a term used in entomology to refer to a form of parental behavior in which an adult (most commonly a hymenopteran such as a bee or wasp) feeds its larvae directly after they have hatched, feeding each larva repeatedly u ...
: a larva's food is supplied gradually as it develops, as is the case in honey bees and some bumblebees.
Solitary and communal bees
Most other bees, including familiar insects such as carpenter bee
Carpenter bees are species in the genus ''Xylocopa'' of the subfamily Xylocopinae. The genus includes some 500 bees in 31 subgenera. The common name "carpenter bee" derives from their nesting behavior; nearly all species burrow into hard plant m ...
s, leafcutter bees
The genus ''Megachile'' is a cosmopolitan group of solitary bees, often called leafcutter bees or leafcutting bees; it also includes the called resin bees and mortar bees. While other genera within the family Megachilidae may chew leaves or pet ...
and mason bees
Mason bee is a name now commonly used for species of bees in the genus ''Osmia'', of the family Megachilidae. Mason bees are named for their habit of using mud or other "masonry" products in constructing their nests, which are made in naturally ...
are solitary in the sense that every female is fertile, and typically inhabits a nest she constructs herself. There is no division of labor so these nests lack queens and ''worker'' bees for these species. Solitary bees typically produce neither honey nor 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 ...
.
Bees collect pollen to feed their young, and have the necessary adaptations to do this. However, certain wasp species such as pollen wasps have similar behaviours, and a few species of bee scavenge
Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a h ...
from carcases to feed their offspring.[ Solitary bees are important pollinators; they gather pollen to provision their nests with food for their brood. Often it is mixed with nectar to form a paste-like consistency. Some solitary bees have advanced types of pollen-carrying structures on their bodies. Very few species of solitary bee are being cultured for commercial pollination. Most of these species belong to a distinct set of ]genera
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
which are commonly known by their nesting behavior or preferences, namely: carpenter bees, sweat bees, mason bees, plasterer bees, 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, dwarf carpenter bees, leafcutter bees, alkali bees and digger bees.
Most solitary bees are fossorial
A fossorial () animal is one adapted to digging which lives primarily, but not solely, underground. Some examples are badgers, naked mole-rats, clams, meerkats, and mole salamanders, as well as many beetles, wasps, and bees.
Prehistoric eviden ...
, digging nests in the ground in a variety of soil textures and conditions, while others create nests in hollow reeds or twigs, or holes in wood
Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin th ...
. The female typically creates a compartment (a "cell") with an egg and some provisions for the resulting larva, then seals it off. A nest may consist of numerous cells. When the nest is in wood, usually the last (those closer to the entrance) contain eggs that will become males. The adult does not provide care for the brood once the egg is laid, and usually dies after making one or more nests. The males typically emerge first and are ready for mating when the females emerge. Solitary bees are very unlikely to sting (only in self-defense, if ever), and some (esp. in the family Andrenidae
The Andrenidae (commonly known as mining bees) are a large, nearly cosmopolitan family of solitary, ground-nesting bees. Most of the family's diversity is located in temperate or arid areas (warm temperate xeric). It includes some enormous gene ...
) are stingless.
While solitary, females each make individual nests. Some species, such as the European mason bee '' Hoplitis anthocopoides'', and the Dawson's Burrowing bee, ''Amegilla dawsoni,'' are gregarious, preferring to make nests near others of the same species, and giving the appearance of being social. Large groups of solitary bee nests are called ''aggregations'', to distinguish them from colonies
In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
. In some species, multiple females share a common nest, but each makes and provisions her own cells independently. This type of group is called "communal" and is not uncommon. The primary advantage appears to be that a nest entrance is easier to defend from predators and parasites when multiple females use that same entrance regularly.[
]
Biology
Life cycle
The life cycle of a bee, be it a solitary or social species, involves the laying of an egg, the development through several moults of a legless 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 ...
, a 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 ...
tion stage during which the insect undergoes complete metamorphosis
Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygo ...
, followed by the emergence of a winged adult. The number of eggs laid by a female during her lifetime can vary from eight or less in some solitary bees, to more than a million in highly social species. Most solitary bees and bumble bees in temperate climates overwinter as adults or pupae and emerge in spring when increasing numbers of flowering plants come into bloom. The males usually emerge first and search for females with which to mate. Like the other members of Hymenoptera bees are haplodiploid
Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky.
Haplodiploidy determines the se ...
; the sex of a bee is determined by whether or not the egg is fertilised. After mating, a female stores the sperm, and determines which sex is required at the time each individual egg is laid, fertilised eggs producing female offspring and unfertilised eggs, males. Tropical bees may have several generations in a year and no diapause
In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
stage.
The egg is generally oblong, slightly curved and tapering at one end. Solitary bees, lay each egg in a separate cell with a supply of mixed pollen and nectar next to it. This may be rolled into a pellet or placed in a pile and is known as mass provisioning. Social bee species provision progressively, that is, they feed the larva regularly while it grows. The nest varies from a hole in the ground or in wood, in solitary bees, to a substantial structure with wax combs in bumblebees and honey bees.
In most species, larvae are whitish grubs, roughly oval and bluntly-pointed at both ends. They have 15 segments and spiracle Spiracle or spiraculum may refer to:
* Spiracle (arthropods), opening in the exoskeletons of some arthropods
* Spiracle (vertebrates), openings on the surface of some vertebrates
* Spiraculum, a genus of land snails in family Cyclophoridae
Cycl ...
s in each segment for breathing. They have no legs but move within the cell, helped by tubercles on their sides. They have short horns on the head, jaws for chewing food and an appendage on either side of the mouth tipped with a bristle. There is a gland under the mouth that secretes a viscous liquid which solidifies into the silk they use to produce a cocoon. The cocoon is semi-transparent and the pupa can be seen through it. Over the course of a few days, the larva undergoes metamorphosis into a winged adult. When ready to emerge, the adult splits its skin dorsally and climbs out of the exuvia
In biology, exuviae are the remains of an exoskeleton and related structures that are left after ecdysozoans (including insects, crustaceans and arachnids) have moulted. The exuviae of an animal can be important to biologists as they can often b ...
e and breaks out of the cell.[
File:Apoidea.jpg, Nest of common carder ]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 ...
, wax canopy removed to show winged 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 pupae
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 ...
in irregularly placed wax cells
File:Carpenter Bee Galleries.jpeg, Carpenter bee
Carpenter bees are species in the genus ''Xylocopa'' of the subfamily Xylocopinae. The genus includes some 500 bees in 31 subgenera. The common name "carpenter bee" derives from their nesting behavior; nearly all species burrow into hard plant m ...
nests in a cedar wood beam (sawn open)
File:Bienen mit Brut 2.jpg, Honeybees on brood comb
Brood may refer to:
Nature
* Brood, a collective term for offspring
* Brooding, the incubation of bird eggs by their parents
* Bee brood, the young of a beehive
* Individual broods of North American Periodical Cicadas:
** Brood X, the largest br ...
with eggs and larvae
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 ...
in cells
Flight
Antoine Magnan
Antoine Magnan (13 June 1881 – 5 March 1938) was a French zoologist and aeronautical engineer who studied the flight of insects and birds for possible lessons to apply to powered flight. He is best known for a remark in his 1934 book ''Le Vol de ...
's 1934 book ''Le vol des insectes'', says that he and André Sainte-Laguë
André Sainte-Laguë (, 20 April 1882 – 18 January 1950) was a French mathematician who was a pioneer in the area of graph theory.
His research on seat allocation methods (published in 1910) led to one being named after him, the Sainte-Laguë ...
had applied the equations of air resistance
In fluid dynamics, drag (sometimes called air resistance, a type of friction, or fluid resistance, another type of friction or fluid friction) is a force acting opposite to the relative motion of any object moving with respect to a surrounding flu ...
to insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s and found that their flight could not be explained by fixed-wing calculations, but that "One shouldn't be surprised that the results of the calculations don't square with reality". This has led to a common misconception that bees "violate aerodynamic theory". In fact it merely confirms that bees do not engage in fixed-wing flight, and that their flight is explained by other mechanics, such as those used by helicopter
A helicopter is a type of rotorcraft in which lift and thrust are supplied by horizontally spinning rotors. This allows the helicopter to take off and land vertically, to hover, and to fly forward, backward and laterally. These attributes ...
s. In 1996 it was shown that vortices created by many insects' wings helped to provide lift. High-speed cinematography
Cinematography (from ancient Greek κίνημα, ''kìnema'' "movement" and γράφειν, ''gràphein'' "to write") is the art of motion picture (and more recently, electronic video camera) photography.
Cinematographers use a lens to focu ...
and robotic mock-up of a bee wing[ Re: work of Dr. Michael H. Dickinson.] showed that lift was generated by "the unconventional combination of short, choppy wing strokes, a rapid rotation of the wing as it flops over and reverses direction, and a very fast wing-beat frequency". Wing-beat frequency normally increases as size decreases, but as the bee's wing beat covers such a small arc, it flaps approximately 230 times per second, faster than a fruitfly (200 times per second) which is 80 times smaller.
Navigation, communication, and finding food
The ethologist Karl von Frisch
Karl Ritter von Frisch, (20 November 1886 – 12 June 1982) was a German-Austrian ethologist who received the Nobel Prize in Physiology or Medicine in 1973, along with Nikolaas Tinbergen and Konrad Lorenz.
His work centered on investigations of ...
studied navigation
Navigation is a field of study that focuses on the process of monitoring and controlling the movement of a craft or vehicle from one place to another.Bowditch, 2003:799. The field of navigation includes four general categories: land navigation, ...
in the honey bee. He showed that honey bees communicate by the waggle 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 nect ...
, in which a worker indicates the location of a food source to other workers in the hive. He demonstrated that bees can recognize a desired compass direction in three different ways: by the sun, by the polarization pattern of the blue sky, and by the earth's magnetic field. He showed that the sun is the preferred or main compass; the other mechanisms are used under cloudy skies or inside a dark beehive
A beehive is an enclosed structure in which some honey bee species of the subgenus '' Apis'' live and raise their young. Though the word ''beehive'' is commonly used to describe the nest of any bee colony, scientific and professional literature ...
. Bees navigate using spatial memory with a "rich, map-like organization".
Digestion
The gut of bees is relatively simple, but multiple metabolic strategies exist in the gut microbiota
Microbiota are the range of microorganisms that may be commensal, symbiotic, or pathogenic found in and on all multicellular organisms, including plants. Microbiota include bacteria, archaea, protists, fungi, and viruses, and have been found t ...
. Pollinating bees consume nectar and pollen, which require different digestion strategies by somewhat specialized bacteria. While nectar is a liquid of mostly monosaccharide
Monosaccharides (from Greek ''monos'': single, '' sacchar'': sugar), also called simple sugars, are the simplest forms of sugar and the most basic units (monomers) from which all carbohydrates are built.
They are usually colorless, water-solub ...
sugars and so easily absorbed, pollen contains complex polysaccharide
Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wa ...
s: branching pectin
Pectin ( grc, πηκτικός ': "congealed" and "curdled") is a heteropolysaccharide, a structural acid contained in the primary lamella, in the middle lamella, and in the cell walls of terrestrial plants. The principal, chemical component of ...
and hemicellulose
A hemicellulose (also known as polyose) is one of a number of heteropolymer, heteropolymers (matrix polysaccharides), such as arabinoxylans, present along with cellulose in almost all embryophyte, terrestrial plant cell walls.Scheller HV, Ulvskov H ...
. Approximately five groups of bacteria are involved in digestion. Three groups specialize in simple sugars ('' Snodgrassella'' and two groups of ''Lactobacillus
''Lactobacillus'' is a genus of Gram-positive, aerotolerant anaerobes or microaerophilic, rod-shaped, non-spore-forming bacteria. Until 2020, the genus ''Lactobacillus'' comprised over 260 phylogenetically, ecologically, and metabolically diver ...
''), and two other groups in complex sugars ('' Gilliamella'' and ''Bifidobacterium
''Bifidobacterium'' is a genus of gram-positive, nonmotile, often branched anaerobic bacteria. They are ubiquitous inhabitants of the gastrointestinal tract though strains have been isolated from the vagina and mouth ('' B. dentium'') of mammal ...
''). Digestion of pectin and hemicellulose is dominated by bacterial 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 ''Gilliamella'' and ''Bifidobacterium
''Bifidobacterium'' is a genus of gram-positive, nonmotile, often branched anaerobic bacteria. They are ubiquitous inhabitants of the gastrointestinal tract though strains have been isolated from the vagina and mouth ('' B. dentium'') of mammal ...
'' respectively. Bacteria that cannot digest polysaccharides obtain enzymes from their neighbors, and bacteria that lack certain amino acids do the same, creating multiple ecological niche
In ecology, a niche is the match of a species to a specific environmental condition.
Three variants of ecological niche are described by
It describes how an organism or population responds to the distribution of resources and competitors (for ...
s.
Although most bee species are nectarivorous
In zoology, a nectarivore is an animal which derives its energy and nutrient requirements from a diet consisting mainly or exclusively of the sugar-rich nectar produced by flowering plants.
Nectar as a food source presents a number of benefits a ...
and palynivorous, some are not. Particularly unusual are vulture bees in the genus ''Trigona
''Trigona'' is one of the largest genera of stingless bees, comprising about 32 species, exclusively occurring in the New World, and formerly including many more subgenera than the present assemblage; many of these former subgenera have been el ...
,'' which consume carrion and wasp brood, turning meat into a honey-like substance.
Ecology
Floral relationships
Most bees are polylectic (generalist) meaning they collect pollen from a range of flowering plants, but some are oligolege
The term oligolecty is used in pollination ecology to refer to bees that exhibit a narrow, specialized preference for pollen sources, typically to a single family or genus of flowering plants. The preference may occasionally extend broadly to mu ...
s (specialists), in that they only gather pollen from one or a few species or genera of closely related plants. Specialist pollinators also include bee species which gather floral oils instead of pollen, and male orchid bees, which gather aromatic compounds from orchid
Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant.
Along with the Asteraceae, they are one of the two largest families of flowering ...
s (one of the few cases where male bees are effective pollinators). Bees are able to sense the presence of desirable flowers through ultraviolet patterning on flowers, floral odors, and even electromagnetic fields. Once landed, a bee then uses nectar quality and pollen taste to determine whether to continue visiting similar flowers.
In rare cases, a plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
species may only be effectively pollinated by a single bee species, and some plants are endangered
An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inva ...
at least in part because their pollinator is also threatened. But, there is a pronounced tendency for oligolectic bees to be associated with common, widespread plants visited by multiple pollinator species. For example, the creosote bush
''Larrea tridentata'', called creosote bush and greasewood as a plant, chaparral as a medicinal herb, and ''gobernadora'' (Spanish language, Spanish for "governess") in Mexico, due to its ability to secure more water by inhibiting the growth of n ...
in the arid parts of the United States southwest is associated with some 40 oligoleges.
As mimics and models
Many bees are aposematic
Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
ally coloured, typically orange and black, warning of their ability to defend themselves with a powerful sting. As such they are models for Batesian mimicry
Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on bu ...
by non-stinging insects such as bee-flies
The Bombyliidae are a family of flies, commonly known as bee flies. Adults generally feed on nectar and pollen, some being important pollinators. Larvae are mostly parasitoids of other insects.
Overview
The Bombyliidae are a large family o ...
, robber flies and hoverflies
Hover flies, also called flower flies or syrphid flies, make up the insect family Syrphidae. As their common name suggests, they are often seen hovering or nectaring at flowers; the adults of many species feed mainly on nectar and pollen, while ...
, all of which gain a measure of protection by superficially looking and behaving like bees.
Bees are themselves Müllerian mimics of other aposematic insects with the same colour scheme, including wasp
A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s, lycid and other beetles, and many butterflies and moths (Lepidoptera
Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
) which are themselves distasteful, often through acquiring bitter and poisonous chemicals from their plant food. All the Müllerian mimics, including bees, benefit from the reduced risk of predation that results from their easily recognised warning coloration.
Bees are also mimicked by plants such as the bee orchid Bee orchid is a common name for several orchids and may refer to:
*'' Cottonia peduncularis'', a species of orchid from India and Sri Lanka
*'' Diuris carinata'', a species of orchid from the south-west of Western Australia
*'' Ida barringtoniae'', ...
which imitates both the appearance and the scent of a female bee; male bees attempt to mate (pseudocopulation
Pseudocopulation describes behaviors similar to copulation that serve a reproductive function for one or both participants but do not involve actual sexual union between the individuals. It is most generally applied to a pollinator attempting to co ...
) with the furry lip of the flower, thus pollinating it.
As brood parasites
Brood parasite
Brood parasites are animals that rely on others to raise their young. The strategy appears among birds, insects and fish. The brood parasite manipulates a host, either of the same or of another species, to raise its young as if it were its own ...
s occur in several bee families including the apid subfamily Nomadinae
Nomadinae is a subfamily of bees in the family Apidae. They are known commonly as cuckoo bees.
This subfamily is entirely kleptoparasitic. They occur worldwide, and use many different types of bees as hosts. As parasites, they lack a pollen-car ...
. Females of these species lack pollen collecting structures (the scopa
''Scopa'' (; literally "broom") is an Italian card game, and one of the three major national card games in Italy, the others being ''Briscola'' and ''Tresette''. It is also popular in Argentina and Brazil, brought in by Italian immigrants, mostly ...
) and do not construct their own nests. They typically enter the nests of pollen collecting species, and lay their eggs in cells provisioned by the host bee. When the "cuckoo" bee larva hatches, it consumes the host larva's pollen ball, and often the host egg also. In particular, the Arctic bee species, ''Bombus hyperboreus
''Bombus hyperboreus'' is a species of Arctic bumblebee with a circumpolar distribution. The species is primarily found in the arctic areas of Greenland, northern Scandinavia, and Russia. In 2015 the nearctic species, '' Bombus natvigi'', was sep ...
'' is an aggressive species that attacks and enslaves other bees of the same subgenus. However, unlike many other bee brood parasites, they have pollen baskets and often collect pollen.
In Southern Africa, hives of African honeybees (''A. mellifera scutellata'') are being destroyed by parasitic workers of the Cape honeybee, ''A. m. capensis''. These lay 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 ...
eggs ("thelytoky
Thelytoky (from the Greek ''thēlys'' "female" and ''tokos'' "birth") is a type of parthenogenesis in which females are produced from unfertilized eggs, as for example in aphids. Thelytokous parthenogenesis is rare among animals and reported in a ...
"), escaping normal worker policing
Worker policing is a behavior seen in colonies of social hymenopterans (ants, bees, and wasps) whereby worker females eat or remove eggs that have been laid by other workers rather than those laid by a queen. Worker policing ensures that the ...
, leading to the colony's destruction; the parasites can then move to other hives.
The cuckoo bee
The term cuckoo bee is used for a variety of different bee lineages which have evolved the kleptoparasitism, kleptoparasitic behaviour of laying their eggs in the nests of other bees, reminiscent of the behavior of cuckoo birds. The name is perhap ...
s in the ''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 ...
'' subgenus ''Psithyrus'' are closely related to, and resemble, their hosts in looks and size. This common pattern gave rise to the ecological principle "Emery's rule
In 1909, the entomologist Carlo Emery noted that social parasites among insects (e.g., kleptoparasites) tend to be parasites of species or genera to which they are closely related.Emery, C. "Über den Ursprung der dulotischen, parasitischen und my ...
". Others parasitize bees in different families, like ''Townsendiella
''Townsendiella'' is a genus of cuckoo bees in the family Apidae, found in Central America, Mexico, and the southwestern United States. It is the sole genus of the tribe Townsendiellini.
Species
* ''Townsendiella californica'' Michener, 1936
* ' ...
'', a nomadine apid, two species of which are cleptoparasites of the dasypodaid genus ''Hesperapis
''Hesperapis'' is a genus of evening bees in the family Melittidae. There are at least 30 described species in ''Hesperapis''.Michez D. (2008) ''Monographic revision of the melittid bees (Hymenoptera, Apoidea, Melittidae sensu lato)].'' Proc. Net ...
'', while the other species in the same genus attacks halictidae, halictid bees.
Nocturnal bees
Four bee families (Andrenidae
The Andrenidae (commonly known as mining bees) are a large, nearly cosmopolitan family of solitary, ground-nesting bees. Most of the family's diversity is located in temperate or arid areas (warm temperate xeric). It includes some enormous gene ...
, Colletidae
The Colletidae are a family of bees, and are often referred to collectively as plasterer bees or polyester bees, due to the method of smoothing the walls of their nest cells with secretions applied with their mouthparts; these secretions dry int ...
, Halictidae
Halictidae is the second-largest family of bees (clade Anthophila) with nearly 4,500 species. Halictid species are an extremely diverse group that can vary greatly in appearance. These bees occur all over the world and are found on every contine ...
, and Apidae
Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bumblebees and honey bees, but also includes stingless bees (also used for ...
) contain some species that are crepuscular
In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
. Most are tropical or subtropical, but some live in arid regions at higher latitudes. These bees have greatly enlarged ocelli
A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
, which are extremely sensitive to light and dark, though incapable of forming images. Some have refracting superposition compound eyes: these combine the output of many elements of their compound eyes to provide enough light for each retinal photoreceptor. Their ability to fly by night enables them to avoid many predators, and to exploit flowers that produce nectar only or also at night.
Predators, parasites and pathogens
Vertebrate predators of bees include bee-eater
The bee-eaters are a group of non-passerine birds in the family Meropidae, containing three genera and thirty species. Most species are found in Africa and Asia, with a few in southern Europe, Australia, and New Guinea. They are characterised by ...
s, shrikes
Shrikes () are passerine birds of the family Laniidae. The family is composed of 34 species in four genera.
The family name, and that of the largest genus, ''Lanius'', is derived from the Latin word for "butcher", and some shrikes are also know ...
and flycatchers, which make short sallies to catch insects in flight. Swifts and swallows fly almost continually, catching insects as they go. The honey buzzard attacks bees' nests and eats the larvae. The greater honeyguide
The greater honeyguide (''Indicator indicator'') is a bird in the family Indicatoridae, paleotropical near passerine birds related to the woodpeckers. Its English and scientific names refer to its habit of guiding people to bee colonies. Claims t ...
interacts with humans by guiding them to the nests of wild bees. The humans break open the nests and take the honey and the bird feeds on the larvae and the wax. Among mammals, predators such as the badger
Badgers are short-legged omnivores in the family Mustelidae (which also includes the otters, wolverines, martens, minks, polecats, weasels, and ferrets). Badgers are a polyphyletic rather than a natural taxonomic grouping, being united b ...
dig up bumblebee nests and eat both the larvae and any stored food.
Specialist ambush predators of visitors to flowers include crab spiders
The Thomisidae are a family of spiders, including about 170 genera and over 2,100 species. The common name crab spider is often linked to species in this family, but is also applied loosely to many other families of spiders. Many members of th ...
, which wait on flowering plants for pollinating insects; predatory bugs, and praying mantis
Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They ha ...
es, some of which (the flower mantis
Flower mantises are praying mantis species that use a special form of camouflage referred to as aggressive mimicry, which they not only use to attract prey, but avoid predators as well. These insects have specific colorations and behaviors that ...
es of the tropics) wait motionless, aggressive mimics camouflaged as flowers. Beewolves are large wasps that habitually attack bees; the ethologist
Ethology is the scientific study of animal behaviour, usually with a focus on behaviour under natural conditions, and viewing behaviour as an evolutionarily adaptive trait. Behaviourism as a term also describes the scientific and objective ...
Niko Tinbergen
Nikolaas "Niko" Tinbergen (; ; 15 April 1907 – 21 December 1988) was a Dutch biologist and ornithologist who shared the 1973 Nobel Prize in Physiology or Medicine with Karl von Frisch and Konrad Lorenz for their discoveries concerning the or ...
estimated that a single colony of the beewolf ''Philanthus triangulum
The European beewolf (''Philanthus triangulum''), also known as the bee-killer wasp or the bee-eating philanthus (from the now obsolete synonym ''Philanthus apivorus''), is a solitary wasp that lives in the Western Palearctic and Afrotropics. Alt ...
'' might kill several thousand honeybees in a day: all the prey he observed were honeybees. Other predatory insects that sometimes catch bees include robber flies and dragonflies
A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
. Honey bees are affected by parasites including tracheal
The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends from the la ...
and ''Varroa
''Varroa'' is a genus of parasitic mesostigmatan mites associated with honey bees, placed in its own family, Varroidae. The genus was named for Marcus Terentius Varro, a Roman scholar and beekeeper. The condition of a honeybee colony being infe ...
'' mite
Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
s. However, some bees are believed to have a mutualistic relationship with mites.
Some mites of genus '' Tarsonemus'' are associated with bees. They live in bee nests and ride on adult bees for dispersal. They are presumed to feed on fungi, nest materials or pollen. However, the impact they have on bees remains uncertain.
Relationship with humans
In mythology and folklore
Homer
Homer (; grc, Ὅμηρος , ''Hómēros'') (born ) was a Greek poet who is credited as the author of the ''Iliad'' and the ''Odyssey'', two epic poems that are foundational works of ancient Greek literature. Homer is considered one of the ...
's '' Hymn to Hermes'' describes three bee-maidens with the power of divination
Divination (from Latin ''divinare'', 'to foresee, to foretell, to predict, to prophesy') is the attempt to gain insight into a question or situation by way of an occultic, standardized process or ritual. Used in various forms throughout histor ...
and thus speaking truth, and identifies the food of the gods as honey. Sources associated the bee maidens with Apollo
Apollo, grc, Ἀπόλλωνος, Apóllōnos, label=genitive , ; , grc-dor, Ἀπέλλων, Apéllōn, ; grc, Ἀπείλων, Apeílōn, label=Arcadocypriot Greek, ; grc-aeo, Ἄπλουν, Áploun, la, Apollō, la, Apollinis, label= ...
and, until the 1980s, scholars followed Gottfried Hermann (1806) in incorrectly identifying the bee-maidens with the Thriae
The Thriae (; grc, Θριαί, Thriaí) were nymphs, three virginal sisters, one of a number of such triads in Greek mythology. They were named ''Melaina'' ("The Black"), ''Kleodora'' ("Famed for her Gift"), and ''Daphnis'' ("Laurel") or ''Cor ...
. Honey, according to a Greek myth, was discovered by a nymph called Melissa ("Bee"); and honey was offered to the Greek gods from Mycenean times. Bees were also associated with the Delphic oracle
Pythia (; grc, Πυθία ) was the name of the high priestess of the Temple of Apollo at Delphi. She specifically served as its oracle and was known as the Oracle of Delphi. Her title was also historically glossed in English as the Pythoness ...
and the prophetess was sometimes called a bee.
The image of a community of honey bees has been used from ancient to modern times, in Aristotle
Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of phil ...
and Plato
Plato ( ; grc-gre, Πλάτων ; 428/427 or 424/423 – 348/347 BC) was a Greek philosopher born in Athens during the Classical period in Ancient Greece. He founded the Platonist school of thought and the Academy, the first institution ...
; in Virgil
Publius Vergilius Maro (; traditional dates 15 October 7021 September 19 BC), usually called Virgil or Vergil ( ) in English, was an ancient Roman poet of the Augustan period. He composed three of the most famous poems in Latin literature: t ...
and Seneca
Seneca may refer to:
People and language
* Seneca (name), a list of people with either the given name or surname
* Seneca people, one of the six Iroquois tribes of North America
** Seneca language, the language of the Seneca people
Places Extrat ...
; in Erasmus
Desiderius Erasmus Roterodamus (; ; English: Erasmus of Rotterdam or Erasmus;''Erasmus'' was his baptismal name, given after St. Erasmus of Formiae. ''Desiderius'' was an adopted additional name, which he used from 1496. The ''Roterodamus'' wa ...
and Shakespeare
William Shakespeare ( 26 April 1564 – 23 April 1616) was an English playwright, poet and actor. He is widely regarded as the greatest writer in the English language and the world's pre-eminent dramatist. He is often called England's nation ...
; Tolstoy
Count Lev Nikolayevich TolstoyTolstoy pronounced his first name as , which corresponds to the romanization ''Lyov''. () (; russian: link=no, Лев Николаевич Толстой,In Tolstoy's day, his name was written as in pre-refor ...
, and by political and social theorists such as Bernard Mandeville
Bernard Mandeville, or Bernard de Mandeville (; 15 November 1670 – 21 January 1733), was an Anglo-Dutch philosopher, political economist and satirist. Born in Rotterdam, Netherlands, he lived most of his life in England and used English for ...
and Karl Marx
Karl Heinrich Marx (; 5 May 1818 – 14 March 1883) was a German philosopher, economist, historian, sociologist, political theorist, journalist, critic of political economy, and socialist revolutionary. His best-known titles are the 1848 ...
as a model for human society
A society is a group of individuals involved in persistent social interaction, or a large social group sharing the same spatial or social territory, typically subject to the same political authority and dominant cultural expectations. Socie ...
. In English folklore, bees would be told of important events in the household, in a custom known as " Telling the bees".
In art and literature
Some of the oldest examples of bees in art are rock paintings
In archaeology, rock art is human-made markings placed on natural surfaces, typically vertical stone surfaces. A high proportion of surviving historic and prehistoric rock art is found in caves or partly enclosed rock shelters; this type also m ...
in Spain
, image_flag = Bandera de España.svg
, image_coat = Escudo de España (mazonado).svg
, national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond")
, national_anthem = (English: "Royal March")
, i ...
which have been dated to 15,000 BC.
W. B. Yeats
William Butler Yeats (13 June 186528 January 1939) was an Irish poet, dramatist, writer and one of the foremost figures of 20th-century literature. He was a driving force behind the Irish Literary Revival and became a pillar of the Irish liter ...
's poem '' The Lake Isle of Innisfree'' (1888) contains the couplet "Nine bean rows will I have there, a hive for the honey bee, / And live alone in the bee loud glade." At the time he was living in Bedford Park in the West of London. Beatrix Potter
Helen Beatrix Potter (, 28 July 186622 December 1943) was an English writer, illustrator, natural scientist, and conservationist. She is best known for her children's books featuring animals, such as ''The Tale of Peter Rabbit'', which was he ...
's illustrated book ''The Tale of Mrs Tittlemouse
''The Tale of Mrs. Tittlemouse'' is a children's book written and illustrated by Beatrix Potter, and published by Frederick Warne & Co. in 1910. The tale is about housekeeping and insect pests in the home, and reflects Potter's own sense of tid ...
'' (1910) features Babbity Bumble and her brood ''(pictured)''. Kit Williams
Christopher "Kit" Williams (born 28 April 1946) is an English artist, illustrator and author best known for his 1979 book '' Masquerade'', a pictorial storybook which contains clues to the location of a golden (18 carat) jewelled hare created ...
' treasure hunt book ''The Bee on the Comb
''The Bee on the Comb'' is an armchair treasure hunt book, written and illustrated by Kit Williams and published in May 1984. It is a follow-up to Williams' previous treasure hunt book '' Masquerade'', although not a direct sequel.
The book wa ...
'' (1984) uses bees and beekeeping as part of its story and puzzle. Sue Monk Kidd
Sue Monk Kidd (born August 12, 1948) is an American writer from Sylvester, Georgia best known for her novels '' The Secret Life of Bees'' and ''The Invention of Wings''.
Early life and education
Kidd was born in Sylvester, Georgia, and attended ...
's '' The Secret Life of Bees'' (2004), and the 2009 film starring Dakota Fanning
Hannah Dakota Fanning (born February 23, 1994) is an American actress. She rose to prominence at the age of seven for her performance as Lucy Dawson in the drama film ''I Am Sam'' (2001), for which she received a Screen Actors Guild Award nomina ...
, tells the story of a girl who escapes her abusive home and finds her way to live with a family of beekeepers, the Boatwrights.
The 2007 animated comedy film ''Bee Movie
''Bee Movie'' is a 2007 American computer-animated comedy film co-produced by DreamWorks Animation SKG and Columbus 81 Productions, and distributed by Paramount Pictures. The film was directed by Simon J. Smith and Steve Hickner (in the f ...
'' used Jerry Seinfeld
Jerome Allen Seinfeld ( ; born April 29, 1954) is an American stand-up comedian, actor, writer, and producer. He is best known for playing a Jerry Seinfeld (character), semi-fictionalized version of himself in the sitcom ''Seinfeld'', which he ...
's first script and was his first work for children; he starred as a bee named Barry B. Benson, alongside Renée Zellweger
Renée Kathleen Zellweger (; born April 25, 1969) is an American actress. The recipient of various accolades, including two Academy Awards, two British Academy Film Awards, and four Golden Globe Awards, she was one of the world's highest-paid ...
. Critics found its premise awkward and its delivery tame. Dave Goulson
Dave Goulson (born 30 July 1965) is Professor of Biology (Evolution, Behaviour and Environment) at the University of Sussex. Specializing in the ecology and conservation of insects, particularly bumblebees, Goulson is the author of several bo ...
's ''A Sting in the Tale
''A Sting in the Tale'' is a 1989 Australian political satire film directed by Eugene Schlusser and starring Diane Craig and Gary Day.David Stratton, ''The Avocado Plantation: Boom and Bust in the Australian Film Industry'', Pan MacMillan, 1990 p ...
'' (2014) describes his efforts to save bumblebees in Britain, as well as much about their biology. The playwright Laline Paull's fantasy
Fantasy is a genre of speculative fiction involving Magic (supernatural), magical elements, typically set in a fictional universe and sometimes inspired by mythology and folklore. Its roots are in oral traditions, which then became fantasy ...
''The Bees'' (2015) tells the tale of a hive bee named Flora 717 from hatching onwards.
Beekeeping
Humans have kept honey bee colonies, commonly in hives
Hives, also known as urticaria, is a kind of skin rash with red, raised, itchy bumps. Hives may burn or sting. The patches of rash may appear on different body parts, with variable duration from minutes to days, and does not leave any long-lasti ...
, for millennia. Beekeeper
A beekeeper is a person who keeps honey bees.
Beekeepers are also called honey farmers, apiarists, or less commonly, apiculturists (both from the Latin '' apis'', bee; cf. apiary). The term beekeeper refers to a person who keeps honey bees i ...
s collect 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 ...
, 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 ...
, propolis
Propolis or bee glue is a resinous mixture that honey bees produce by mixing saliva and beeswax with exudate gathered from tree buds, sap flows, or other botanical sources. It is used as a sealant for unwanted open spaces in the beehive. Pro ...
, 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 ...
, and 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 ...
from hives; bees are also kept to pollinate
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, a ...
crops and to produce bees for sale to other beekeepers.
Depictions of humans collecting honey from wild bees date to 15,000 years ago; efforts to domesticate them are shown in Egyptian art around 4,500 years ago. Simple hives and smoke were used; jars of honey were found in the tombs of pharaoh
Pharaoh (, ; Egyptian: ''pr ꜥꜣ''; cop, , Pǝrro; Biblical Hebrew: ''Parʿō'') is the vernacular term often used by modern authors for the kings of ancient Egypt who ruled as monarchs from the First Dynasty (c. 3150 BC) until the an ...
s such as Tutankhamun
Tutankhamun (, egy, twt-ꜥnḫ-jmn), Egyptological pronunciation Tutankhamen () (), sometimes referred to as King Tut, was an Egyptian pharaoh who was the last of his royal family to rule during the end of the Eighteenth Dynasty (ruled ...
. From the 18th century, European understanding of the colonies and biology of bees allowed the construction of the moveable comb hive so that honey could be harvested without destroying the colony. Among Classical Era authors, beekeeping with the use of smoke is described in Aristotle's ''History of Animals
''History of Animals'' ( grc-gre, Τῶν περὶ τὰ ζῷα ἱστοριῶν, ''Ton peri ta zoia historion'', "Inquiries on Animals"; la, Historia Animalium, "History of Animals") is one of the major texts on biology by the ancient Gr ...
'' Book 9. The account mentions that bees die after stinging; that workers remove corpses from the hive, and guard it; castes including workers and non-working drone
Drone most commonly refers to:
* Drone (bee), a male bee, from an unfertilized egg
* Unmanned aerial vehicle
* Unmanned surface vehicle, watercraft
* Unmanned underwater vehicle or underwater drone
Drone, drones or The Drones may also refer to:
...
s, but "kings" rather than queens; predators including toads and bee-eaters; and the waggle 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 nect ...
, with the "irresistible suggestion" of άροσειονται ("aroseiontai", it waggles) and παρακολουθούσιν ("parakolouthousin", they watch).
Beekeeping is described in detail by Virgil
Publius Vergilius Maro (; traditional dates 15 October 7021 September 19 BC), usually called Virgil or Vergil ( ) in English, was an ancient Roman poet of the Augustan period. He composed three of the most famous poems in Latin literature: t ...
in his ''Georgics
The ''Georgics'' ( ; ) is a poem by Latin poet Virgil, likely published in 29 BCE. As the name suggests (from the Greek word , ''geōrgika'', i.e. "agricultural (things)") the subject of the poem is agriculture; but far from being an example ...
''; it is also mentioned in his ''Aeneid
The ''Aeneid'' ( ; la, Aenē̆is or ) is a Latin Epic poetry, epic poem, written by Virgil between 29 and 19 BC, that tells the legendary story of Aeneas, a Troy, Trojan who fled the Trojan_War#Sack_of_Troy, fall of Troy and travelled to ...
'', and in Pliny's '' Natural History''.[
]
As commercial pollinators
Bees play an important role in pollinating
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, a ...
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s, and are the major type of pollinator
A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains.
Insects are the maj ...
in many ecosystem
An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
s that contain flowering plants. It is estimated that one third of the human food supply depends on 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 ...
by insects, birds and bats, most of which is accomplished by bees, whether wild or domesticated. Over the last half century, there has been a general decline in the species richness of wild bees and other pollinators, probably attributable to stress from increased parasites and disease, the use of pesticides, and a general decrease in the number of wild flowers. Climate change probably exacerbates the problem.
Contract pollination has overtaken the role of honey production for beekeeper
A beekeeper is a person who keeps honey bees.
Beekeepers are also called honey farmers, apiarists, or less commonly, apiculturists (both from the Latin '' apis'', bee; cf. apiary). The term beekeeper refers to a person who keeps honey bees i ...
s in many countries. After the introduction of Varroa mites
''Varroa destructor'', the ''Varroa'' mite is an external parasitic mite that attacks and feeds on the honey bees ''Apis cerana'' and ''Apis mellifera''. The disease caused by the mites is called varroosis.
The ''Varroa'' mite can reproduce on ...
, 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 ...
honey bees declined dramatically in the US, though their numbers have since recovered. The number of colonies kept by beekeepers declined slightly, through urbanization
Urbanization (or urbanisation) refers to the population shift from rural to urban areas, the corresponding decrease in the proportion of people living in rural areas, and the ways in which societies adapt to this change. It is predominantly t ...
, systematic pesticide use, tracheal
The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends from the la ...
and ''Varroa
''Varroa'' is a genus of parasitic mesostigmatan mites associated with honey bees, placed in its own family, Varroidae. The genus was named for Marcus Terentius Varro, a Roman scholar and beekeeper. The condition of a honeybee colony being infe ...
'' mites, and the closure of beekeeping businesses. In 2006 and 2007 the rate of attrition increased, and was described as colony collapse disorder
Colony collapse disorder (CCD) is an abnormal phenomenon that occurs when the majority of worker bees in a honey bee colony disappear, leaving behind a queen, plenty of food, and a few nurse bees to care for the remaining immature bees. While s ...
. In 2010 invertebrate iridescent virus and the fungus ''Nosema ceranae
''Nosema ceranae'' is a microsporidian, a small, unicellular parasite that mainly affects ''Apis cerana'', the Asiatic honey bee. Along with '' Nosema apis'', it causes the disease nosemosis, the most widespread of the diseases of adult honey b ...
'' were shown to be in every killed colony, and deadly in combination. Winter losses increased to about 1/3. ''Varroa'' mites were thought to be responsible for about half the losses.
Apart from colony collapse disorder, losses outside the US have been attributed to causes including pesticide seed dressings, using 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 ...
s such as Clothianidin
Clothianidin is an insecticide developed by Takeda Chemical Industries and Bayer AG. Similar to thiamethoxam and imidacloprid, it is a neonicotinoid. Neonicotinoids are a class of insecticides that are chemically similar to nicotine, which has be ...
, Imidacloprid
Imidacloprid is a systemic insecticide belonging to a class of chemicals called the neonicotinoids which act on the central nervous system of insects. The chemical works by interfering with the transmission of stimuli in the insect nervous system. ...
and Thiamethoxam
Thiamethoxam is the ISO common name for a mixture of ''cis-trans'' isomers used as a systemic insecticide of the neonicotinoid class. It has a broad spectrum of activity against many types of insects and can be used as a seed dressing.
Histor ...
. From 2013 the European Union
The European Union (EU) is a supranational political and economic union of member states that are located primarily in Europe. The union has a total area of and an estimated total population of about 447million. The EU has often been des ...
restricted some pesticides to stop bee populations from declining further. In 2014 the Intergovernmental Panel on Climate Change
The Intergovernmental Panel on Climate Change (IPCC) is an intergovernmental body of the United Nations. Its job is to advance scientific knowledge about climate change caused by human activities. The World Meteorological Organization (WMO) a ...
report warned that bees faced increased risk of extinction because of global warming
In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...
. In 2018 the European Union decided to ban field use of all three major neonicotinoids; they remain permitted in veterinary, greenhouse, and vehicle transport usage.
Farmers have focused on alternative solutions to mitigate these problems. By raising native plants, they provide food for native bee pollinators like ''Lasioglossum vierecki
''Lasioglossum vierecki'', also known as ''Dialictus vierecki'' and ''Halictus vierecki'',various contributors. 2015. Hymenoptera Online (HOL). nlineAvailable fro ccessed 22 September 2015/ref> is a sand sweat bee and is part of the family Hal ...
''[Kuehn, F. Coordinator. (2015). Farming for native bees. World Wide Web electronic publication. Retrieved from . (Accessed: 22 September 2015).] and '' L. leucozonium'',[Adamson, Nancy Lee]
An Assessment of Non-Apis Bees as Fruit and Vegetable Crop Pollinators in Southwest Virginia
. Diss. 2011. Web. 15 October 2015. leading to less reliance on honey bee populations.
File:Peponapis pruinosaCane-12.JPG, 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 (Apidae) are important pollinators of squashes and cucumber
Cucumber (''Cucumis sativus'') is a widely-cultivated Vine#Horticultural climbing plants, creeping vine plant in the Cucurbitaceae family that bears usually cylindrical Fruit, fruits, which are used as culinary vegetables. s.
File:A bee covered with pollen.jpg, Bee covered in pollen
As food producers
Honey is a natural product produced by bees and stored for their own use, but its sweetness has always appealed to humans. Before domestication of bees was even attempted, humans were raiding their nests for their honey. Smoke was often used to subdue the bees and such activities are depicted in rock paintings
In archaeology, rock art is human-made markings placed on natural surfaces, typically vertical stone surfaces. A high proportion of surviving historic and prehistoric rock art is found in caves or partly enclosed rock shelters; this type also m ...
in Spain dated to 15,000 BC.[
Honey bees are used commercially to produce ]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 ...
. They also produce some substances used as dietary supplements with possible health benefits, pollen, propolis
Propolis or bee glue is a resinous mixture that honey bees produce by mixing saliva and beeswax with exudate gathered from tree buds, sap flows, or other botanical sources. It is used as a sealant for unwanted open spaces in the beehive. Pro ...
, and 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 ...
, though all of these can also cause allergic reactions
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 ...
.
As food
Bees are considered edible insects
Insects as food or edible insects are insect species used for human consumption. More than 2,000 insects species worldwide are considered edible. However, a much smaller number is discussed for industrialized mass production and partly regiona ...
. People in some countries eat insects, including the larvae and pupae of bees, mostly stingless species. They also gather larvae, pupae and surrounding cells, known as bee brood
In beekeeping, bee brood or brood refers to the eggs, larvae and pupae of honey bee, honeybees. The brood of Western honey bees develops within a Beehive (beekeeping), bee hive. In man-made, removable frame hives, such as Langstroth hives, each f ...
, for consumption. In 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 dish '' botok tawon'' from Central and East Java
Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's List ...
, bee larvae are eaten as a companion to 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 ...
, after being mixed with shredded coconut
The coconut tree (''Cocos nucifera'') is a member of the palm tree family ( Arecaceae) and the only living species of the genus ''Cocos''. The term "coconut" (or the archaic "cocoanut") can refer to the whole coconut palm, the seed, or the ...
, wrapped in banana
A banana is an elongated, edible fruit – botanically a berry – produced by several kinds of large herbaceous flowering plants in the genus ''Musa''. In some countries, bananas used for cooking may be called "plantains", distinguis ...
leaves, and steamed.[ (This particular Botok recipe uses anchovies, not bees)]
Bee brood (pupae and larvae) although low in calcium
Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to ...
, has been found to be high in 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 ...
and carbohydrate
In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ma ...
, and a useful source of phosphorus
Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
, magnesium
Magnesium is a chemical element with the symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 of the periodic ta ...
, potassium
Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosphe ...
, and trace minerals iron
Iron () is a chemical element with symbol Fe (from la, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, right in f ...
, zinc
Zinc is a chemical element with the symbol Zn and atomic number 30. Zinc is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodi ...
, copper
Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...
, and selenium
Selenium is a chemical element with the symbol Se and atomic number 34. It is a nonmetal (more rarely considered a metalloid) with properties that are intermediate between the elements above and below in the periodic table, sulfur and tellurium, ...
. In addition, while bee brood was high in fat, it contained no fat soluble vitamins (such as A, D, and E) but it was a good source of most of the water-soluble B-vitamins
B vitamins are a class of water-soluble vitamins that play important roles in cell metabolism and synthesis of red blood cells. Though these vitamins share similar names (B1, B2, B3, etc.), they are chemically distinct compounds that often coexist ...
including choline Choline is an essential nutrient for humans and many other animals. Choline occurs as a cation that forms various salts (X− in the depicted formula is an undefined counteranion). Humans are capable of some ''de novo synthesis'' of choline but re ...
as well as vitamin C. The fat was composed mostly of saturated
Saturation, saturated, unsaturation or unsaturated may refer to:
Chemistry
* Saturation, a property of organic compounds referring to carbon-carbon bonds
** Saturated and unsaturated compounds
**Degree of unsaturation
** Saturated fat or fatty ac ...
and monounsaturated fatty acids with 2.0% being polyunsaturated fatty acids
Polyunsaturated fatty acids (PUFAs) are fatty acids that contain more than one double bond in their backbone. This class includes many important compounds, such as essential fatty acids and those that give drying oils their characteristic proper ...
.
File:Botoktawon.jpg, Bee larvae as food in the Java
Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's List ...
nese dish '' botok tawon''
File:Fried bees dish.jpg, Fried whole bees served in a Ukrainian restaurant
As alternative medicine
Apitherapy
Apitherapy is a branch of alternative medicine that uses honey bee products, including honey, pollen, propolis, royal jelly and bee venom. There has been no scientific or clinical evidence for the efficacy or safety of apitherapy treatments.
...
is a branch of alternative medicine
Alternative medicine is any practice that aims to achieve the healing effects of medicine despite lacking biological plausibility, testability, repeatability, or evidence from clinical trials. Complementary medicine (CM), complementary and alt ...
that uses honey bee products, including raw honey, 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 ...
, pollen, propolis
Propolis or bee glue is a resinous mixture that honey bees produce by mixing saliva and beeswax with exudate gathered from tree buds, sap flows, or other botanical sources. It is used as a sealant for unwanted open spaces in the beehive. Pro ...
, 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 ...
and apitoxin (Bee venom
Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a sti ...
). The claim that apitherapy treats cancer, which some proponents of apitherapy make, remains unsupported by evidence-based medicine
Evidence-based medicine (EBM) is "the conscientious, explicit and judicious use of current best evidence in making decisions about the care of individual patients". The aim of EBM is to integrate the experience of the clinician, the values of t ...
.
Stings
The painful stings of bees are mostly associated with the poison gland and the Dufour's gland
Dufour's gland is an abdominal gland of certain insects, part of the anatomy of the ovipositor or sting apparatus in female members of Apocrita. The diversification of Hymenoptera took place in the Cretaceous and the gland may have developed at a ...
which are abdominal exocrine glands containing various chemicals. In ''Lasioglossum leucozonium
''Lasioglossum leucozonium'' (Schrank, 1781), also known as ''Lasioglossum similis'', is a widespread solitary sweat bee found in North America, Europe, Asia, and parts of northern Africa. While now a common bee in North America, population gen ...
'', the Dufour's Gland mostly contains octadecanolide as well as some eicosanolide. There is also evidence of n-triscosane, n-heptacosane
Higher alkanes are alkanes having nine or more carbon atoms. Nonane is the lightest alkane to have a flash point above 25 °C, and is not classified as dangerously flammable.
The term ''higher alkanes'' is sometimes used literally as "alka ...
, and 22-docosanolide. However, the secretions of these glands could also be used for nest construction.
See also
* Australian native bees
Australia has over 1,700 species of native bee. Bees collect pollen from flowers to feed their young. Flies do not do this, although they may be seen ''eating'' pollen, so identification is not always easy.
Sting or no sting, solitary vs ...
* Superorganism
A superorganism or supraorganism is a group of synergetically interacting organisms of the same species. A community of synergetically interacting organisms of different species is called a holobiont.
Concept
The term superorganism is used mo ...
* World Bee Day
World Bee Day is celebrated on May 20. On this day Anton Janša, the pioneer of beekeeping, was born in 1734.
The purpose of the international day is to acknowledge the role of bees and other pollinators for the ecosystem.
The UN Member Stat ...
Explanatory notes
References
External links
*
"Apoidea"
at All Living Thingsimages, identification guides, and maps of bees
Anthophila (Apoidea) – Bees
orth American species of bees at BugGuide
BugGuide (or BugGuide.net) is a website and online community of naturalists, both amateur and professional, who share observations of arthropods such as insects, spiders, and other related creatures. The website consists of informational guide p ...
Native Bees of North America
at BugGuide
BugGuide (or BugGuide.net) is a website and online community of naturalists, both amateur and professional, who share observations of arthropods such as insects, spiders, and other related creatures. The website consists of informational guide p ...
"Bee declines driven by combined stress from parasites, pesticides, and lack of flowers"
'Science
Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and predictions about the universe.
Science may be as old as the human species, and some of the earliest archeological evidence for ...
''
{{Authority control
Extant Early Cretaceous first appearances