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59-497: Clade Anthophila The superfamily Apoidea is a major group within the Hymenoptera , which includes two traditionally recognized lineages, the "sphecoid" wasps , and the bees . Molecular phylogeny demonstrates that the bees arose from within the traditional " Crabronidae ", so that grouping is paraphyletic , and this has led to a reclassification to produce monophyletic families. Bees appear in recent classifications to be

118-500: A brain and a ventral nerve cord . Most insects reproduce by laying eggs . Insects breathe air through a system of paired openings along their sides, connected to small tubes that take air directly to the tissues. The blood therefore does not carry oxygen; it is only partly contained in vessels, and some circulates in an open hemocoel . Insect vision is mainly through their compound eyes , with additional small ocelli . Many insects can hear, using tympanal organs , which may be on

177-488: A clade , called Anthophila . There are over 20,000 known species of bees in seven recognized biological families . Some species – including honey bees , bumblebees , and stingless bees  – live socially in colonies while most species (>90%) – including mason bees , carpenter bees , leafcutter bees , and sweat bees  – are solitary. Bees are found on every continent except Antarctica , in every habitat on

236-401: A division of labour into reproductive and non-reproductive adults, plus overlapping generations. 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,

295-663: 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 pupation stage during which the insect undergoes complete metamorphosis , 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

354-429: 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

413-466: A distinct set of genera which are commonly known by their nesting behavior or preferences, namely: carpenter bees, sweat bees , mason bees, plasterer bees , squash bees , dwarf carpenter bees , leafcutter bees, alkali bees and digger bees . Most solitary bees are fossorial , 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 . The female typically creates

472-563: A hundred species, are marine. Insects such as snow scorpionflies flourish in cold habitats including the Arctic and at high altitude. Insects such as desert locusts , ants, beetles, and termites are adapted to some of the hottest and driest environments on earth, such as the Sonoran Desert . Insects form a clade , a natural group with a common ancestor, among the arthropods . A phylogenetic analysis by Kjer et al. (2016) places

531-525: A larva's food is supplied gradually as it develops, as is the case in honey bees and some bumblebees. Most other bees, including familiar insects such as carpenter bees , leafcutter bees and mason bees 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 . Bees collect pollen to feed their young, and have

590-440: 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

649-729: A male is haploid (has only one copy of each gene), his daughters (which are diploid , 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 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

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708-625: A nearly immobile pupa . Insects that undergo three-stage metamorphosis lack a pupa, developing through a series of increasingly adult-like nymphal stages. The higher level relationship of the insects is unclear. Fossilized insects of enormous size have been found from the Paleozoic Era, including giant dragonfly-like insects with wingspans of 55 to 70 cm (22 to 28 in). The most diverse insect groups appear to have coevolved with flowering plants . Adult insects typically move about by walking and flying; some can swim. Insects are

767-478: A paraphyletic "Crabronidae." Ampulicidae (Cockroach wasps) [REDACTED] Astatidae [REDACTED] Bembicidae [REDACTED] Sphecidae ( sensu stricto ) [REDACTED] Crabronidae ( sensu stricto ) [REDACTED] Mellinidae [REDACTED] Pemphredonidae [REDACTED] Philanthidae [REDACTED] Psenidae [REDACTED] Ammoplanidae Anthophila (bees) [REDACTED] This Apoidea -related article

826-533: 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 . 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 bees (Apidae) include some primitively eusocial species with similar biology. Some allodapine bees (Apidae) form primitively eusocial colonies, with progressive provisioning :

885-399: A specialized lineage of "crabronid" wasps that switched to the use of pollen and nectar as larval food, rather than insect prey; this makes the traditional "Crabronidae" a paraphyletic group. Accordingly, bees and sphecoids are now all grouped together in a single superfamily, and the older available name is "Apoidea" rather than "Sphecoidea" (which, like Spheciformes, has been used in

944-525: A variety of ways. Male moths can sense the pheromones of female moths over great distances. Other species communicate with sounds: crickets stridulate , or rub their wings together, to attract a mate and repel other males. Lampyrid beetles communicate with light. Humans regard many insects as pests , especially those that damage crops, and attempt to control them using insecticides and other techniques. Others are parasitic , and may act as vectors of diseases . Insect pollinators are essential to

1003-512: Is Cretotrigona prisca , a corbiculate bee of Late Cretaceous age (~70 mya) found in New Jersey amber . 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", 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

1062-407: Is a stub . You can help Misplaced Pages by expanding it . Bee Apiformes (from Latin 'apis' ) 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 superfamily Apoidea . They are currently considered

1121-690: Is based on Sann et al. , 2018, which used phylogenomics to demonstrate that both the bees ( Anthophila ) and the Sphecidae arose from within the former "Crabronidae," which is therefore paraphyletic , and which they suggested should be split into several families; the former family Heterogynaidae nests in Nyssonini within the Bembicidae , as defined by these authors. These findings differ in several details from studies published by two other sets of authors in 2017, though all three studies demonstrate

1180-476: Is believed that mites eat fungi that attack pollen, so the relationship in this case may be mutualistic . Molecular phylogeny was used by Debevic et al , 2012, to demonstrate that the bees ( Anthophila ) arose from deep within the Crabronidae sensu lato , which was thus rendered paraphyletic . In their study, the placement of the monogeneric Heterogynaidae was uncertain. The small family Mellinidae

1239-444: 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. 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. The most advanced of these are species with eusocial colonies; these are characterised by cooperative brood care and

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1298-555: 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. The oldest non-compression bee fossil

1357-541: 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. Genomic analysis indicates that despite only appearing much later in

1416-853: The Latin word insectum from in , "cut up", as insects appear to be cut into three parts. The Latin word was introduced by Pliny the Elder who calqued the Ancient Greek word ἔντομον éntomon "insect" (as in entomology ) from ἔντομος éntomos "cut in pieces"; this was Aristotle 's term for this class of life in his biology , also in reference to their notched bodies. The English word insect first appears in 1601 in Philemon Holland 's translation of Pliny. In common speech, insects and other terrestrial arthropods are often called bugs . Entomologists to some extent reserve

1475-707: The Stenotritidae ), and by the end of the Cretaceous, South American bees had also colonized North America. 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 . Following the K-Pg extinction, surviving bee lineages continued to spread into

1534-439: 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 of jointed legs , compound eyes , and a pair of antennae . Insects are the most diverse group of animals, with more than a million described species ; they represent more than half of all animal species. The insect nervous system consists of

1593-588: The Bumblebee Specialist Group to review the threat status of all bumblebee species worldwide using the IUCN Red List 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 , 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

1652-597: 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. 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

1711-623: The Hemiptera (true bugs), Lepidoptera (butterflies and moths), Diptera (true flies), Hymenoptera (wasps, ants, and bees), and Coleoptera (beetles), each with more than 100,000 described species. Insects are distributed over every continent and almost every terrestrial habitat. There are many more species in the tropics , especially in rainforests , than in temperate zones. The world's regions have received widely differing amounts of attention from entomologists. The British Isles have been thoroughly surveyed, so that Gullan and Cranston 2014 state that

1770-592: 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. A second extinction event among bees

1829-488: The appearance of being social. Large groups of solitary bee nests are called aggregations , to distinguish them from colonies . 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. The life cycle of

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1888-848: 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. Melittidae (inc. Dasypodainae , Meganomiinae ) at least 50 Mya [REDACTED] Apidae (inc. honeybees, cuckoo bees, carpenter bees) ≈87 Mya [REDACTED] Megachilidae (mason, leafcutter bees) ≈50 Mya [REDACTED] Andrenidae (mining bees) ≈34 Mya [REDACTED] Halictidae (sweat bees) ≈50 Mya [REDACTED] Colletidae (plasterer bees) ≈25 Mya [REDACTED] Stenotritidae (large Australian bees) ≈2 Mya [REDACTED] Bees differ from closely related groups such as wasps by having branched or plume-like setae (hairs), combs on

1947-677: 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 , native to Europe, the Middle East, and Africa. Africanized bees are a hybrid strain of A. mellifera that escaped from experiments involving crossing European and African subspecies; they are extremely defensive. Stingless bees are also highly eusocial . They practise mass provisioning , with complex nest architecture and perennial colonies also established via swarming. Many bumblebees are eusocial, similar to

2006-401: 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 ( meliponiculture for stingless bees) has been practised for millennia, since at least

2065-487: The eusocial Vespidae such as hornets 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 set up

2124-480: 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

2183-530: The females emerge. Solitary bees are very unlikely to sting (only in self-defense, if ever), and some (esp. in the family Andrenidae ) 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

2242-605: 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

2301-470: 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: 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

2360-460: 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,

2419-464: 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. True honey bees (genus Apis , of which eight species are currently recognized) are highly eusocial, and are among

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2478-608: The insects among the Hexapoda , six-legged animals with segmented bodies; their closest relatives are the Diplura (bristletails). Collembola (springtails) [REDACTED] Protura (coneheads) [REDACTED] Diplura (two-pronged bristletails) [REDACTED] Insecta (=Ectognatha) [REDACTED] The internal phylogeny is based on the works of Wipfler et al. 2019 for the Polyneoptera , Johnson et al. 2018 for

2537-660: 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

2596-406: The legs or other parts of the body. Their sense of smell is via receptors, usually on the antennae and the mouthparts. Nearly all insects hatch from eggs . Insect growth is constrained by the inelastic exoskeleton, so development involves a series of molts . The immature stages often differ from the adults in structure, habit and habitat. Groups that undergo four-stage metamorphosis often have

2655-661: The lineage as incertae sedis within the Aculeata . The Allodapini (within the Apidae) appeared 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

2714-442: The multiple (at least nine) evolutions of eusociality within Hymenoptera . 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. But, monogamy (queens mating singly)

2773-402: The name "bugs" for a narrow category of " true bugs ", insects of the order Hemiptera , such as cicadas and shield bugs . Other terrestrial arthropods, such as centipedes , millipedes , woodlice , spiders , mites and scorpions , are sometimes confused with insects, since they have a jointed exoskeleton. Adult insects are the only arthropods that ever have wings, with up to two pairs on

2832-565: The necessary adaptations to do this. However, certain wasp species such as pollen wasps have similar behaviours, and a few species of bee scavenge 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

2891-554: The only invertebrates that can achieve sustained powered flight; insect flight evolved just once. Many insects are at least partly aquatic , and have larvae with gills; in some species, the adults too are aquatic. Some species, such as water striders , can walk on the surface of water. Insects are mostly solitary, but some, such as bees , ants and termites , are social and live in large, well-organized colonies . Others, such as earwigs , provide maternal care, guarding their eggs and young. Insects can communicate with each other in

2950-495: The other members of Hymenoptera bees are haplodiploid ; 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 stage. Insect Insects (from Latin insectum ) are hexapod invertebrates of

3009-643: The past, but also defined a paraphyletic group and has been abandoned). As bees (not including their wasp ancestors) are still considered a monophyletic group, they are given a grouping between superfamily and family to unify all bees, Anthophila. Chrysidoidea (cuckoo wasps and allies) Vespoidea (potter wasps, paper wasps, and others) Sierolomorphidae (sierolomorphid wasps) Tiphiidae (tiphiid wasps) Thynnoidea (thynnid wasps and others) Pompiloidea (spider wasps, velvet ants, and others) Scolioidea (scoliid wasps) Formicoidea (ants) Apoidea (spheciform wasps and bees) This phylogenetic tree

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3068-547: 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 the leafcutter bee Megachile pluto , the largest species of bee, whose females can attain

3127-535: The reproduction of many flowering plants and so to their ecosystems. Many insects are ecologically beneficial as predators of pest insects, while a few provide direct economic benefit. Two species in particular are economically important and were domesticated many centuries ago: silkworms for silk and honey bees for honey . Insects are consumed as food in 80% of the world's nations, by people in roughly 3000 ethnic groups. Human activities are having serious effects on insect biodiversity . The word insect comes from

3186-527: The thorax. Whether winged or not, adult insects can be distinguished by their three-part body plan, with head, thorax, and abdomen; they have three pairs of legs on the thorax. Estimates of the total number of insect species vary considerably, suggesting that there are perhaps some 5.5 million insect species in existence, of which about one million have been described and named. These constitute around half of all eukaryote species, including animals , plants , and fungi . The most diverse insect orders are

3245-492: 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. The immediate ancestors of bees were stinging wasps in

3304-525: The total of around 22,500 species is probably within 5% of the actual number there; they comment that Canada's list of 30,000 described species is surely over half of the actual total. They add that the 3000 species of the American Arctic must be broadly accurate. In contrast, a large majority of the insect species of the tropics and the southern hemisphere are probably undescribed. Some 30–40,000 species inhabit freshwater ; very few insects, perhaps

3363-604: The tribe Meliponini whose workers are less than 2 millimetres (0.08 in) in length. 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. In haplodiploid species, females develop from fertilized eggs and males from unfertilized eggs. Because

3422-501: 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

3481-839: Was not included in this analysis. Further studies by Sann et al. , 2018, elevated the subfamilies (plus one tribe and one subtribe) of Crabronidae sensu lato to family status. They also recovered the placement of Heterogyna within Nyssonini and sunk Heterogynaidae . The newly erected family, Ammoplanidae , formerly a subtribe of Pemphredoninae , was recovered as the most sister family to bees. Ampulicidae (Cockroach wasps) [REDACTED] Astatidae [REDACTED] Bembicidae [REDACTED] Sphecidae ( sensu stricto ) [REDACTED] Crabronidae ( sensu stricto ) [REDACTED] Mellinidae [REDACTED] Pemphredonidae [REDACTED] Philanthidae [REDACTED] Psenidae [REDACTED] Ammoplanidae Anthophila (bees) [REDACTED] This cladogram of

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