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There are over 20,000 bee species worldwide, including the honey bee, which originated in Eurasia and has been imported around the globe as a domesticated species. Wild bees species live on every continent except Antarctica. In North America there are approximately 4,000 native bee species occupying ecosystems from forests to deserts to grasslands.

Though all female bees can sting, they only do so when threatened. Honey bees, with hives filled with honey and larvae that need protecting, are generally more aggressive and likely to sting when disturbed than solitary native bees.

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Queen bumble bees can live for a year and workers for a month. Solitary bees also live for about a year, with the majority of that time spent developing in their nesting chamber where they hatch, pupate, and often overwinter. Their adult lives, during which they are active, lasts approximately three to eight weeks. Females tend to live a bit longer, as they need to build a nest and lay eggs.

Both domesticated honey bees and many native bee species are in decline. In fact some species, such as the once-common rusty patched bumblebee, are now listed as endangered in the U.S. Potential causes of these declines include habitat destruction, disease, agricultural and lawn and garden practices, use of pesticides, habitat fragmentation, changes in land use, invasive species, and climate change.

Bees are found on every continent except 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,[5] 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 primates and 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.[6]

Human beekeeping or apiculture (meliponiculture for stingless bees) 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. In Mesoamerica, the Mayans have practiced large-scale intensive meliponiculture since pre-Columbian times.[5]

The immediate ancestors of bees were stinging 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.

Based on phylogenetic analysis, bees are thought to have originated during the Early Cretaceous (about 124 million years ago) on the supercontinent of West Gondwana, just prior to its breakup into South America and Africa. The supercontinent is thought to have been a largely xeric environment at this time; modern bee diversity hotspots are also in xeric and seasonal temperate environments, suggesting strong niche conservatism among bees ever since their origins.[7]

Genomic analysis indicates that despite only appearing much later in the fossil record, all modern bee families had already diverged from one another by the end of the Cretaceous. The Melittidae, Apidae, and Megachilidae had already evolved on the supercontinent prior to its fragmentation. Further divergences were facilitated by West Gondwana's breakup around 100 million years ago, leading to a deep Africa-South America split within both the Apidae and Megachilidae, the isolation of the Melittidae in Africa, and the origins of the Colletidae, Andrenidae and Halictidae in South America. The rapid radiation of the South American bee families is thought to have followed the concurrent radiation of flowering plants in the same region. Later in the Cretaceous (80 million years ago), colletid bees colonized Australia from South America (with an offshoot lineage evolving into the Stenotritidae), and by the end of the Cretaceous, South American bees had also colonized North America.[7] The North American fossil taxon Cretotrigona belongs to a group that is no longer found in North America, suggesting that many bee lineages went extinct during the Cretaceous-Paleogene extinction event.[7]

Following the K-Pg extinction, surviving bee lineages continued to spread into the Northern Hemisphere, colonizing Europe from Africa by the Paleocene, and then spreading east to Asia. This was facilitated by the warming climate around the same time, allowing bees to move to higher latitudes following the spread of tropical and subtropical habitats. By the Eocene (~45 mya) there was already considerable diversity among eusocial bee lineages.[8][a] A second extinction event among bees is thought to have occurred due to rapid climatic cooling around the Eocene-Oligocene boundary, leading to the extinction of some bee lineages such as the tribe Melikertini. Over the Paleogene and Neogene, different bee lineages continued to spread all over the world, and the shifting habitats and connectedness of continents led to the isolation and evolution of many new bee tribes.[7]

The oldest non-compression bee fossil is Cretotrigona prisca, a corbiculate bee of Late Cretaceous age (~70 mya) found in New Jersey amber.[11] A fossil from the early Cretaceous (~100 mya), Melittosphex burmensis, was initially considered "an extinct lineage of pollen-collecting Apoidea sister to the modern bees",[12] but subsequent research has rejected the claim that Melittosphex is a bee, or even a member of the superfamily Apoidea to which bees belong, instead treating the lineage as incertae sedis within the Aculeata.[13]

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[23] such as nectar and longer tubes, and bees developed longer tongues to extract the nectar.[24] 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.[3] The appearance of these structures drove the adaptive radiation of the angiosperms, and, in turn, bees themselves.[9] 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 mutualistic.[25][26]

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.[27] The small subfamily Mellininae was not included in this analysis.

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).[31] The smallest species may be dwarf stingless bees in the tribe Meliponini whose workers are less than 2 millimetres (0.08 in) in length.[32]

According to inclusive fitness 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 species such as bees because of their unusual relatedness structure.[33]

Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such as termites 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.[37] 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.[35]

Bees may be solitary or may live in various types of communities. Eusociality appears to have originated from at least three independent origins in halictid bees.[38] The most advanced of these are species with eusocial colonies; these are characterised by cooperative brood care and a division of labour into reproductive and non-reproductive adults, plus overlapping generations.[39] This division of labour creates specialized groups within eusocial societies which are called castes. 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. The group is called eusocial if, in addition, the group consists of a mother (the queen) and her daughters (workers). When the castes are purely behavioural alternatives, with no morphological differentiation other than size, the system is considered primitively eusocial, as in many paper wasps; when the castes are morphologically discrete, the system is considered highly eusocial.[24] ff782bc1db