Bees are flying insects closely related to wasps and ants, known for their role 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 superfamily Apoidea. They are presently considered a clade, called Anthophila. There are over 16,000 known species of bees in seven recognized biological families.

Some species — including honey beesbumblebees, and stingless bees — live socially in colonies while some species — including mason bees, carpenter bees, leafcutter bees, and sweat bees — are solitary.

Bees are found on every continent except for Antarctica, in every habitat on the planet that contains insect-pollinated flowering plants.

The most common bees in the Northern Hemisphere are the Halictidae, or sweat bees, but they are small and often mistaken for wasps or flies. Bees range in size from tiny stingless bee species, whose workers are less than 2 millimetres (0.08 in) long, to Megachile pluto, the largest species of leafcutter bee, whose females can attain a length of 39 millimetres (1.54 in).

Bees feed on nectar and pollen, the former primarily as an energy source and the latter primarily for protein and other nutrients.

Most pollen is used as food for their larvae. Vertebrate predators of bees include birds such as bee-eaters; insect predators include beewolves and dragonflies.

Bee pollination is important both ecologically and commercially, 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 or apiculture has been practised for millennia, since at least the times of Ancient Egypt and Ancient Greece.

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 where beekeeping is far more common.

Evolution

Melittosphex burmensis, a fossil bee preserved in amber from the Early Cretaceous (100 to 145 million years ago) of Myanmar

The ancestors of bees were wasps in the family Crabronidae, which were predators of other insects. The switch from insect prey to pollen may have resulted from the consumption of prey insects which were flower visitors and were partially covered with pollen when they were fed to the wasp larvae.

This same evolutionary scenario may have occurred within the vespoid wasps, where the pollen wasps evolved from predatory ancestors. Until recently, the oldest non-compression bee fossil had been found in New Jersey amber, Cretotrigona prisca of Cretaceous age, a corbiculate bee.

A bee fossil from the early Cretaceous (~100 mya), Melittosphex burmensis, is considered “an extinct lineage of pollen-collecting Apoidea sister to the modern bees”.

Derived features of its morphology (apomorphies) place it clearly within the bees, but it retains two unmodified ancestral traits (plesiomorphies) of the legs (two mid-tibial spurs, and a slender hind basitarsus), showing its transitional status.

By the Eocene (~45 mya) there was already considerable diversity among eusocial bee lineages.

The highly eusocial corbiculate Apidae appeared roughly 87 Mya, and the Allodapini (within the Apidae) around 53 Mya.

The Colletidae appear as fossils only from the late Oligocene (~25 Mya) to early Miocene. The Melittidae are known from Palaeomacropis eocenicus in the Early Eocene.

The Megachilidae are known from trace fossils (characteristic leaf cuttings) from the Middle Eocene.

The Andrenidae are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale.

The Halictidae first appear in the Early Eocene with species found in amber. The Stenotritidae are known from fossil brood cells of Pleistocene age.

Coevolution

Long-tongued bees and long-tubed flowers coevolved, like this Amegilla cingulata (Apidae) on Acanthus ilicifolius.

The earliest animal-pollinated flowers were shallow, cup-shaped blooms pollinated by insects such as beetles, 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, flowers developed floral rewards such as nectar and longer tubes, and bees developed longer tongues to extract the nectar.

Bees also developed structures known as scopal hairs and pollen baskets 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 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 of the angiosperms, and, in turn, bees themselves.

Bees coevolved 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 mutualistc.

Phylogeny

External

This phylogenetic tree is based on Debevic et al, 2012, which used molecular phylogeny to demonstrate that the bees (Anthophila) arose from deep within the Crabronidae, which is therefore paraphyletic. The placement of the Heterogynaidae is uncertain. The small subfamily Mellininae 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

The lapping mouthparts of a honey bee, showing labium and maxillae

Bees differ from closely related groups such as wasps by having branched or plume-like setae (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 which cover much of the surface of the head. Between and above these are three small simple eyes (ocelli) 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 and a long proboscis 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.

 

Head-on view of a male carpenter bee, showing antennae, three ocelli, compound eyes, and mouthparts

The largest species of bee is thought to be Wallace’s giant bee Megachile pluto, whose females can attain a length of 39 millimetres (1.54 in).

The smallest species may be dwarf stingless bees in the tribe Meliponini whose workers are less than 2 millimetres (0.08 in) in length.

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