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105-547: A honey bee (also spelled honeybee ) is a eusocial flying insect within the genus Apis of the bee clade , all native to mainland Afro-Eurasia . After bees spread naturally throughout Africa and Eurasia , humans became responsible for the current cosmopolitan distribution of honey bees, introducing multiple subspecies into South America (early 16th century ), North America (early 17th century), and Australia (early 19th century). Honey bees are known for their construction of perennial colonial nests from wax ,
210-478: A multicellular organism ; castes fulfill a specific role that contributes to the functioning and survival of the whole colony, while being incapable of independent survival outside the colony. In 1969, Charles D. Michener further expanded Batra's classification with his comparative study of social behavior in bees. He observed multiple species of bees ( Apoidea ) in order to investigate the different levels of animal sociality, all of which are different stages that
315-697: A pest in some regions. However, these strains do not overwinter well, so they are not often found in the colder, more northern parts of North America. The original breeding experiment for which the East African lowland honey bees were brought to Brazil in the first place has continued (though not as originally intended). Novel hybrid strains of domestic and re-domesticated Africanized honey bees combine high resilience to tropical conditions and good yields. They are popular among beekeepers in Brazil. Tribe Apini Latreille Genus Apis Linnaeus ( sensu lato ) As in
420-429: A colony may pass through. Eusociality, which is the highest level of animal sociality a species can attain, specifically had three characteristics that distinguished it from the other levels: E. O. Wilson extended the terminology to include other social insects, such as ants, wasps, and termites. Originally, it was defined to include organisms (only invertebrates) that had the following three features: Eusociality
525-407: A colony, there are both pleated fan-shaped "nest" fronds that collect and hold water, and gutter-shaped "strap" fronds that channel water: no solitary Platycerium species has both types. At the bottom of a colony, there are "nest" fronds that clasp the trunk of the tree supporting the fern, and drooping photosynthetic fronds. These are argued to be adapted to support the colony structurally, i.e. that
630-920: A diverse combination of pollen and nectar. Pollen is the only natural protein source for honey bees. Adult worker honey bees consume 3.4–4.3 mg of pollen per day to meet a dry matter requirement of 66–74% protein. The rearing of one larva requires 125-187.5 mg pollen or 25–37.5 mg protein for proper development. Dietary proteins are broken down into amino acids, ten of which are considered essential to honey bees: methionine, tryptophan, arginine, lysine, histidine, phenylalanine, isoleucine, threonine, leucine, and valine. Of these amino acids, honey bees require highest concentrations of leucine, isoleucine, and valine, however elevated concentrations of arginine and lysine are required for brood rearing. In addition to these amino acids, some B vitamins including biotin, folic acid, nicotinamide, riboflavin, thiamine, pantothenate, and most importantly, pyridoxine are required to rear larvae. Pyridoxine
735-401: A few other types of eusocial bees, a colony generally contains one queen bee , a female; seasonally up to a few thousand drone bees , or males; and tens of thousands of female worker bees . Details vary among the different species of honey bees, but common features include: In cold climates, honey bees stop flying when the temperature drops below about 10 °C (50 °F) and crowd into
840-684: A given area means that the pollen they carry for any one species is often very diluted. As such, they can provide some pollination to many plants, but most plants have some native pollinator that is more effective at pollinating that species. When honey bees are present as an invasive species in an area, they compete for flowers with native pollinators, which can actually push out the native species. Western honey bees have been described as essential to human food production, leading to claims that without their pollination humanity would starve or die out. Apples, blueberries, and cherries, for example, are 90 percent dependent on honeybee pollination. Albert Einstein
945-419: A hive of half-sisters that share only 25% of their genes. The association between haplodiploidy and eusociality is below statistical significance. Haplodiploidy is thus neither necessary nor sufficient for eusociality to emerge. Relatedness does still play a part, as monogamy (queens mating singly) is the ancestral state for all eusocial species so far investigated. If kin selection is an important force driving
1050-407: A human being to death if provoked. Eastern Apis species include three or four species, including A. koschevnikovi , A. nigrocincta , and A. cerana . The genetics of the western honey bee ( A. mellifera ) are unclear. Koschevnikov's honey bee ( Apis koschevnikovi ) is often referred to in the literature as the "red bee of Sabah"; however, A. koschevnikovi
1155-648: A male is haploid, his daughters 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 first termed "supersisters", more closely related to their sisters than they would be to their own offspring. Even though workers often do not reproduce, they can pass on more of their genes by helping to raise their sisters than by having their own offspring (each of which would only have 50% of their genes). This unusual situation, where females may have greater fitness when they help rear sisters rather than producing offspring,
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#17327655099561260-508: A pheromone, helping their nest mates to find the food. Reproductive specialization generally involves the production of sterile members of the species, which carry out specialized tasks to care for the reproductive members. Individuals may have behavior and morphology modified for group defense, including self-sacrificing behavior . For example, members of the sterile caste of the honeypot ants such as Myrmecocystus fill their abdomens with liquid food until they become immobile and hang from
1365-466: A possible resolution to the paradox might lie in the close family relationship, which W.D. Hamilton quantified a century later with his 1964 inclusive fitness theory. After the gene-centered view of evolution was developed in the mid-1970s, non-reproductive individuals were seen as an extended phenotype of the genes, which are the primary beneficiaries of natural selection. According to inclusive fitness theory, organisms can gain fitness by increasing
1470-581: A set of pheromones that alter the behavior of specific castes in the colony. These pheromones may act across different species, as observed in Apis andreniformis (black dwarf honey bee), where worker bees responded to queen pheromone from the related Apis florea (red dwarf honey bee). Pheromones are sometimes used in these castes to assist with foraging. Workers of the Australian stingless bee Tetragonula carbonaria , for instance, mark food sources with
1575-587: A single 14 million-year-old specimen from Nevada. The close relatives of modern honey bees – e.g., bumblebees and stingless bees – are also social to some degree, and social behavior is considered to be a trait that predates the origin of the genus. Among the extant members of Apis , the more basal species make single, exposed combs, while the more recently evolved species nest in cavities and have multiple combs, which has greatly facilitated their domestication. While about 20,000 species of bees exist, only eight species of honey bee are recognized, with
1680-416: A species. Multilevel selection has been heavily criticized for its conflict with the kin selection theory. A reversal to solitarity is an evolutionary phenomenon in which descendants of a eusocial group evolve solitary behavior once again. Bees have been model organisms for the study of reversal to solitarity, because of the diversity of their social systems. Each of the four origins of eusociality in bees
1785-594: A total of 43 subspecies , although historically seven to 11 species are recognized: Apis andreniformis (the black dwarf honey bee); Apis cerana (the eastern honey bee); Apis dorsata (the giant honey bee); Apis florea (the red dwarf honey bee); Apis koschevnikovi (Koschevnikov's honey bee); Apis laboriosa (the Himalayan giant honey bee); Apis mellifera (the western honey bee); and Apis nigrocincta (the Philippine honey bee). Honey bees are
1890-488: A weak form of eusociality. It has been suggested that the colonial and epiphytic staghorn fern , too, may make use of a primitively eusocial division of labor. The term "eusocial" was introduced in 1966 by Suzanne Batra , who used it to describe nesting behavior in Halictid bees, on a scale of subsocial/solitary, colonial/communal, semisocial, and eusocial, where a colony is started by a single individual. Batra observed
1995-529: Is 50%. Therefore, helping behavior is only advantageous if it is biased to helping sisters, which would drive the population to a 1:3 sex ratio of males to females. At this ratio, males, as the rarer sex, increase in reproductive value, reducing the benefit of female-biased investment. Further, not all eusocial species are haplodiploid: termites, some snapping shrimps, and mole rats are not. Conversely, many non-eusocial bees are haplodiploid, and among eusocial species many queens mate with multiple males, resulting in
2100-752: Is a kind or type of bee. It is incorrect to run the two words together, as in dragonfly or butterfly , which are appropriate because dragonflies and butterflies are not flies. Honey bee, not honeybee, is the listed common name in the Integrated Taxonomic Information System , the Entomological Society of America Common Names of Insects Database, and the Tree of Life Web Project . Honey bees appear to have their center of origin in South and Southeast Asia (including
2205-446: Is a species of ambrosia beetle native to Australia, and is the first beetle (order Coleoptera ) to be recognized as eusocial. This species forms colonies in which a single female is fertilized, and is protected by many unfertilized females, which serve as workers excavating tunnels in trees. This species has cooperative brood care, in which individuals care for juveniles that are not their own. Some gall-inducing insects , including
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#17327655099562310-427: Is an aggregation of relatives (because the shrimp do not have to disperse to find food), and much competition for those nesting sites. Being the target of attack promotes a good defense system (soldier caste); soldiers promote the fitness of the whole nest by ensuring safety and reproduction of the queen. Eusociality offers a competitive advantage in shrimp populations. Eusocial species are more abundant, occupy more of
2415-617: Is commonly measured in the billions of dollars, credited with adding about 9% to the value of crops across the world. However, despite contributing substantially to crop pollination, there is debate about the potential spillover to natural landscapes and competition between managed honey bees and many of the ~20,000 species of wild pollinators. Species of Apis are generalist floral visitors, and pollinate many species of flowering plants, but because of their "generalized" nature, they often do so inefficiently. Without specialized adaptations for specific flowers, their ability to reach pollen and nectar
2520-501: Is correlated with the number of females in the nest, implying that having more females leads to more efficient building and provisioning of cells. In similar species with only one queen, such as Lasioglossum malachurum in Europe, the degree of eusociality depends on the clime in which the species is found. Termites (order Blattodea , infraorder Isoptera ) make up another large portion of highly advanced eusocial animals. The colony
2625-563: Is differentiated into various castes: the queen and king are the sole reproducing individuals; workers forage and maintain food and resources; and soldiers defend the colony against ant attacks. The latter two castes, which are sterile and perform highly specialized, complex social behaviors, are derived from different stages of pluripotent larvae produced by the reproductive caste. Some soldiers have jaws so enlarged (specialized for defense and attack) that they are unable to feed themselves and must be fed by workers. Austroplatypus incompertus
2730-615: Is distinguished from all other social systems because individuals of at least one caste usually lose the ability to perform behaviors characteristic of individuals in another caste. Eusocial colonies can be viewed as superorganisms . Eusociality has evolved among the insects , crustaceans , trematoda and mammals . It is most widespread in the Hymenoptera ( ants , bees , and wasps ) and in Blattodea ( termites ). A colony has caste differences: queens and reproductive males take
2835-526: Is limited, foragers will collect water from streams or ponds to meet the needs of the hive. The only domesticated species of honey bee are A. mellifera and A. cerana , and they are often maintained, fed, and transported by beekeepers. In Japan, where A. mellifera is vulnerable to local hornets and disease, the Japanese honey bee A. cerana japonica is used in its place. Modern hives also enable beekeepers to transport bees, moving from field to field as
2940-542: Is not divided between castes. Though controversial, it has been suggested that male homosexuality and female menopause could have evolved through kin selection . This would mean that humans sometimes exhibit a type of alloparental behavior known as " helpers at the nest ", with juveniles and sexually mature adolescents helping their parents raise subsequent broods, as in some birds, some non-eusocial bees , and meerkats . These species are not eusocial: they do not have castes, and helpers reproduce on their own if given
3045-538: Is now Nevada during the Middle Miocene period. It was discovered at Stewart Valley, Nevada, and described by Michael S. Engel , Ismael A. Hinojosa-Díaz, and Alexandr P. Rasnitsyn in 2009. The species belongs to the extinct armbrusteri species group, is most similar to the extinct species Apis armbrusteri from the Miocene of southwestern Germany, and both species may be related by geological range. This
3150-497: Is often invoked to explain the multiple independent evolutions of eusociality (at least nine separate times) within the Hymenoptera. Against the supposed benefits of haplodiploidy for eusociality, Robert Trivers notes that while females share 75% of genes with their sisters in haplodiploid populations, they only share 25% of their genes with their brothers. Accordingly, the average relatedness of an individual to their sibling
3255-400: Is often limited. This combined with their behavioural flexibility may be why they are the most commonly documented pollen thieves . Indeed, for plant species with more specialized pollinators, experiments show that increased honeybee visitation can actually reduce pollination, both where honey bees are non-native and even where they are native. What's more, their tendency to visit all species in
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3360-687: Is often reliably predicted by ecological conditions, and switches in behavioral type have been experimentally induced by translocating offspring of solitary or social populations to warm and cool climates. In H. rubicundus , females produce a single brood in cooler regions and two or more broods in warmer regions, so the former populations are solitary while the latter are social. In another species of sweat bees, L. calceatum , social phenotype has been predicted by altitude and micro-habitat composition, with social nests found in warmer, sunnier sites, and solitary nests found in adjacent, cooler, shaded locations. Facultatively social bee species, however, which comprise
3465-651: Is pale reddish in Sabah State , Borneo , Malaysia , but a dark, coppery colour in the Malay Peninsula and Sumatra , Indonesia . Its habitat is limited to the tropical evergreen forests of the Malay Peninsula , Borneo and Sumatra and they do not live in tropical evergreen rain forests which extend into Thailand , Myanmar , Cambodia and Vietnam . Apis nigrocincta is a cavity-nesting species. The species has rust-coloured scapes , legs, and clypeuses , with reddish-tan hair colour that covers most of
3570-468: Is probably the product of a largely Early Pleistocene radiation aided by climate and habitat changes during the last ice age . That the western honey bee has been intensively managed by humans for many millennia – including hybridization and introductions – has apparently increased the speed of its evolution and confounded the DNA sequence data to a point where little of substance can be said about
3675-670: Is sometimes misquoted as saying "If bees disappeared off the face of the earth, man would only have four years left to live" . Einstein did not say this and there is no science to support this prediction. Many important crops need no insect pollination at all. The ten most important crops, comprising 60% of all human food energy, fall into this category: plantains are sterile and propagated by cuttings, as are cassava ; potatoes , yams , and sweet potatoes are root vegetables propagated by tubers ; soybeans are self-pollinated ; and rice , wheat , sorghum , and maize , are wind-pollinated , as are most other grasses. No crops originating in
3780-430: Is the highest level of organization of sociality . It is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generations within a colony of adults , and a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society, sometimes called castes. Eusociality
3885-640: Is the most prevalent B vitamin found in royal jelly and concentrations vary throughout the foraging season with lowest concentrations found in May and highest concentrations found in July and August. Honey bees lacking dietary pyridoxine were unable to rear brood. Pollen is also a lipid source for honey bees ranging from 0.8% to 18.9%. Lipids are metabolized during the brood stage for precursors required for future biosynthesis. Fat-soluble vitamins A, D, E, and K are not considered essential but have shown to significantly improve
3990-497: The Damaraland mole-rat ( Fukomys damarensis ), both of which are highly inbred . Usually living in harsh or limiting environments, these mole-rats aid in raising siblings and relatives born to a single reproductive queen. However, this classification is controversial owing to disputed definitions of 'eusociality'. To avoid inbreeding , mole rats sometimes outbreed and establish new colonies when resources are sufficient. Most of
4095-534: The Late Miocene . This would fit the hypothesis that the ancestral stock of cave-nesting honey bees was separated into the western group of East Africa and the eastern group of tropical Asia by desertification in the Middle East and adjacent regions, which caused declines of food plants and trees that provided nest sites, eventually causing gene flow to cease. The diversity of A. mellifera subspecies
4200-465: The New World depend on the western honey bee ( Apis mellifera ) at all, as the bee is an invasive species brought over with colonists in the last few centuries. Tomatoes , peppers , squash , and all other New World crops evolved with native pollinators such as squash bees , bumble bees , and other native bees. The stingless bees mentioned by Jefferson are distant relatives of the honey bees, in
4305-626: The Philippines ), as all the extant species except Apis mellifera are native to that region. Notably, living representatives of the earliest lineages to diverge ( Apis florea and Apis andreniformis ) have their center of origin there. The first Apis bees appear in the fossil record at the Eocene – Oligocene boundary (34 mya ), in European deposits. The origin of these prehistoric honey bees does not necessarily indicate Europe as
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4410-619: The Tian Shan range. It is variously called the European, western, or common honey bee in different parts of the world. Many subspecies have adapted to the local geographic and climatic environments; in addition, breeds such as the Buckfast bee have been bred. Behavior, colour, and anatomy can be quite different from one subspecies or even strain to another. A. mellifera phylogeny is the most enigmatic of all honey bee species. It seems to have diverged from its eastern relatives only during
4515-478: The University of Maryland and Auburn University published in 2023 found the number of United States honeybee colonies "remained relatively stable" although 48% of colonies were lost in the year that ended April 1, 2023, with a 12-year average annual mortality rate of 39.6%. The previous year (2021-2022) the loss was 39% and the 2020-2021 loss was 50.8%. Beekeepers told the surveying scientists that 21% loss over
4620-471: The gall -forming aphid , Pemphigus spyrothecae (order Hemiptera ), and thrips such as Kladothrips (order Thysanoptera ), are described as eusocial. These species have very high relatedness among individuals due to their asexual reproduction (sterile soldier castes being clones produced by parthenogenesis ), but the gall-inhabiting behavior gives these species a defensible resource. They produce soldier castes for fortress defense and protection of
4725-519: The gene-centered view of evolution , eusociality was seen as paradoxical: if adaptive evolution unfolds by differential reproduction of individual organisms, the evolution of individuals incapable of passing on their genes presents a challenge. In On the Origin of Species , Darwin referred to the existence of sterile castes as the "one special difficulty, which at first appeared to me insuperable, and actually fatal to my theory". Darwin anticipated that
4830-584: The stingless bees belonging to the genus Melipona and the Indian stingless or dammar bee Tetragonula iridipennis . Modern humans also use beeswax in making candles , soap , lip balms and various cosmetics , as a lubricant and in mould-making using the lost wax process . The genus name Apis is Latin for "bee". Although modern dictionaries may refer to Apis as either honey bee or honeybee, entomologist Robert Snodgrass asserts that correct usage requires two words, i.e., honey bee , because it
4935-485: The western honey bee as well as Apis cerana , are known to engage in effective methods of nest thermoregulation during periods of varying temperature in both summer and winter. During the summer, however, this is achieved through fanning and water evaporation from water collected in various fields. Of all the honey bee species, only A. mellifera has been used extensively for commercial pollination of fruit and vegetable crops. The scale of these pollination services
5040-601: The Philippines to satisfaction; some researchers argue that these are indeed distinct species, but that A. cerana as defined is still paraphyletic , consisting of several separate species, though other researchers argue cerana is a single monophyletic species. A. mellifera , the most common domesticated species, was first domesticated before 2600 BC and was the third insect to have its genome mapped. It seems to have originated in eastern tropical Africa and spread from there to Europe and eastwards into Asia to
5145-470: The United States, and a number of species of Bombus (bumblebees) for pollination in various regions globally, such as tomatoes , which are not effectively pollinated by honey bees. Primarily in places where western honey bees were imported by humans, periodic collapses in western honey bee populations have occurred at least since the late 19th century. However, as humans continued to manipulate
5250-424: The anatomy of the worker caste, which is sterile and provides support for the reproductive caste. Most eusocial societies exist in arthropods , while a few are found in mammals . Some ferns may exhibit a primitive form of eusocial behavior. Eusociality has evolved multiple times in different insect orders, including hymenopterans, termites, thrips, aphids, and beetles. The order Hymenoptera contains
5355-509: The body. Apis cerana , the eastern honey bee proper, is the traditional honey bee of southern and eastern Asia. One of its subspecies, the Indian honey bee ( A. c. indica ), was domesticated and kept in hives in a fashion similar to A. mellifera , though on a more limited, regional scale. It has not been possible yet to resolve its relationship to the Bornean honey bee A. c. nuluensis and Apis nigrocincta from
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#17327655099565460-430: The bristles (setae) guarding the spiracles are firm enough to prevent the mites' entry into the tracheae. Mite infestations are known as acarine, and have been called "Isle of Wight disease". Larval stages of the moth Galleria mellonella parasitize both wild and cultivated honey bees, in particular Apis mellifera and Apis cerana . Eggs are laid within the hive, and the larvae that hatch tunnel through and destroy
5565-585: The ceilings of the underground nests, acting as food storage for the rest of the colony. Not all social insects have distinct morphological differences between castes. For example, in the Neotropical social wasp Synoeca surinama , caste ranks are determined by social displays in the developing brood. These castes are sometimes further specialized in their behavior based on age, as in Scaptotrigona postica workers. Between approximately 0–40 days old,
5670-428: The central area of the hive to form a "winter cluster". The worker bees huddle around the queen bee at the center of the cluster, shivering to keep the center between 27 °C (81 °F) at the start of winter (during the broodless period) and 34 °C (93 °F) once the queen resumes laying. The worker bees rotate through the cluster from the outside to the inside so that no bee gets too cold. The outside edges of
5775-403: The cluster stay at about 8–9 °C (46–48 °F). The colder the weather is outside, the more compact the cluster becomes. During winter, they consume their stored honey to produce body heat. The amount of honey consumed during the winter is a function of winter length and severity, but ranges in temperate climates from 15 to 50 kilograms (33 to 110 lb). In addition, certain bees, including
5880-421: The colony against predators, kleptoparasites , and competitors. In these groups, eusociality is produced by both high relatedness and by living in a restricted, shared area. Eusociality has evolved in three different lineages in the colonial crustacean genus Synalphaeus . S. regalis , S. microneptunus , S. filidigitus , S. elizabethae , S. chacei , S. riosi , S. duffyi , and S. cayoneptunus are
5985-523: The colony, it is protected by a specialized caste of sterile soldier trematodes. Soldiers are smaller, more mobile, and develop along a different pathway than sexually mature reproductives. One difference is that a soldier's mouthparts (pharynx) is five times as big as those of the reproductives. They make up nearly a quarter of the volume of the soldier. These soldiers do not have a germinal mass, never metamorphose to be reproductive, and are, therefore, obligately sterile. Soldiers are readily distinguished from
6090-432: The cooperative behavior of the bees, males and females alike, as they took responsibility for at least one duty (i.e., burrowing, cell construction, oviposition ) within the colony. The cooperativeness was essential as the activity of one labor division greatly influenced the activity of another. Eusocial colonies can be viewed as superorganisms , with individual castes being analogous to different tissue or cell types in
6195-422: The cost of investing in non-reproductive offspring, and an increased risk of disease. All reversals to solitarity have occurred among primitively eusocial groups; none have followed the emergence of advanced eusociality. The "point of no return" hypothesis posits that the morphological differentiation of reproductive and non-reproductive castes prevents highly eusocial species such as the honeybee from reverting to
6300-500: The crop needs pollinating and allowing the beekeeper to charge for the pollination services they provide, revising the historical role of the self-employed beekeeper, and favoring large-scale commercial operations. Bees of various types other than honey bees are also domesticated and used for pollination or other means around the world, including Tetragonula iridipennis in India, the blue orchard bee for tree nut and fruit pollination in
6405-588: The crops that depend on western honey bees for pollination have also been imported since colonial times. Escaped swarms (known as "wild" honey bees, but actually feral ) spread rapidly as far as the Great Plains , usually preceding the colonists. Honey bees did not naturally cross the Rocky Mountains ; they were transported by the Mormon pioneers to Utah in the late 1840s, and by ship to California in
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#17327655099566510-680: The dwarf mongoose, the breeding pair receives food priority and protection from subordinates and rarely has to defend against predators. Scientists have debated whether humans are prosocial or eusocial. Edward O. Wilson called humans eusocial apes, arguing for similarities to ants, and observing that early hominins cooperated to rear their children while other members of the same group hunted and foraged. Wilson and others argued that through cooperation and teamwork, ants and humans form superorganisms. Wilson's claims were vigorously rejected by critics of group selection theory, which grounded Wilson's argument, and because human reproductive labor
6615-491: The early 1850s. Africanized honey bees (known colloquially as "killer bees") are hybrids between European stock and the East African lowland subspecies A. m. scutellata . They are often more aggressive than European honey bees and do not create as much of a honey surplus, but are more resistant to disease and are better foragers. Accidentally released from quarantine in Brazil , they have spread to North America and constitute
6720-438: The early phase of infection. This strategic positioning allows them to effectively defend against invaders, similar to how soldier distribution patterns are seen in other animals with defensive castes. They "appear to be an obligately sterile physical caste, akin to that of the most advanced social insects". Among mammals, two species in the rodent group Phiomorpha are eusocial, the naked mole-rat ( Heterocephalus glaber ) and
6825-509: The eight recorded species of parasitic shrimp that rely on fortress defense and live in groups of closely related individuals in tropical reefs and sponges. They live eusocially with a single breeding female, and a large number of male defenders armed with enlarged snapping claws. There is a single shared living space for the colony members, and the non-breeding members act to defend it. The fortress defense hypothesis additionally points out that because sponges provide both food and shelter, there
6930-524: The eusocial phenotype, with one queen supported by sterile workers and sexually active males ( drones ). In queenless colonies, the lack of queen pheromones causes winged females to quickly shed their wings, develop ovaries and lay eggs. These virgin replacement queens assume the role of the queen and start to produce queen pheromones. Similarly, queen weaver ants Oecophylla longinoda have exocrine glands that produce pheromones which prevent workers from laying reproductive eggs. Similar mechanisms exist in
7035-424: The eusocial wasp Vespula vulgaris . For a queen to dominate all the workers, usually numbering more than 3000 in a colony, she signals her dominance with pheromones. The workers regularly lick the queen while feeding her, and the air-borne pheromone from the queen's body alerts those workers of her dominance. Apis nearctica Apis nearctica is an extinct species of honey bee which existed in what
7140-722: The evolution of eusociality is supported by evidence such as experimentally induced reproductive division of labor, for example when normally solitary queens are forced together. Conversely, female Damaraland mole-rats undergo hormonal changes that promote dispersal after periods of high rainfall. Climate too appears to be a selective agent driving social complexity; across bee lineages and Hymenoptera in general, higher forms of sociality are more likely to occur in tropical than temperate environments. Similarly, social transitions within halictid bees , where eusociality has been gained and lost multiple times, are correlated with periods of climatic warming. Social behavior in facultative social bees
7245-437: The evolution of eusociality, monogamy should be the ancestral state, because it maximizes the relatedness of colony members. Increased parasitism and predation rates are the primary ecological drivers of social organization. Group living affords colony members defense against enemies, specifically predators, parasites, and competitors, and allows them to gain advantage from superior foraging methods. The importance of ecology in
7350-622: The exact relationships of many A. mellifera subspecies. Apis mellifera is not native to the Americas , so it was not present when the European explorers and colonists arrived. However, other native bee species were kept and traded by indigenous peoples. In 1622, European colonists brought the German honey bee ( A. m. mellifera ) to the Americas first, followed later by the Italian honey bee ( A. m. ligustica ) and others. Many of
7455-480: The first decade of the 21st century, abnormally high die-offs (30–70% of hives) of western honey bee colonies have occurred in North America. This has been dubbed "colony collapse disorder" (CCD) and was at first unexplained. It seems to be caused by a combination of factors rather than a single pathogen or poison , possibly including neonicotinoid pesticides or Israeli acute paralysis virus . A survey by
7560-529: The genus Melipona . Still, honey bees are considered "crucial to the food supply, pollinating more than 100 of the crops we eat, including nuts, vegetables, berries, citrus and melons." The USDA reports "Three-fourths of the world’s flowering plants and about 35 percent of the world’s food crops depend on animal pollinators to reproduce" and honey bees "pollinate 80 percent of all flowering plants, including more than 130 types of fruits and vegetables." Honey bees obtain all of their nutritional requirements from
7665-449: The habitat, and use more of the available resources than non-eusocial species. The trematodes are a class of parasitic flatworm, also known as flukes. One species, Haplorchis pumilio , has evolved eusociality involving a colony creating a class of sterile soldiers. One fluke invades a host and establishes a colony of dozens to thousands of clones that work together to take it over. Since rival trematode species can invade and replace
7770-469: The honeycombs that contain bee larva and their honey stores. The tunnels they create are lined with silk, which entangles and starves emerging bees. Destruction of honeycombs also result in honey leaking and being wasted. Both G. mellonella adults and larvae are possible vectors for pathogens that can infect bees, including the Israeli acute paralysis virus and the black queen cell virus . To manage
7875-429: The hormonally based behavioral development of workers and suppress their ovarian development. Both behavioral effects mediated by the nervous system often leading to recognition of queens ( releaser ) and physiological effects on the reproductive and endocrine system ( primer ) are attributed to the same pheromones. These pheromones volatilize or are deactivated within thirty minutes, allowing workers to respond rapidly to
7980-416: The immature and mature reproductive worms. Soldiers are more aggressive than reproductives, attacking heterospecific trematodes that infect their host in vitro . H. pumilio soldiers do not attack conspecifics from other colonies. The soldiers are not evenly distributed throughout the host body. They are found in the highest numbers in the basal visceral mass, where competing trematodes tend to multiply during
8085-558: The individuals cooperatively care for the brood of a single reproductive female (the queen) to which they are most likely related. Thus, it is uncertain whether mole rats are truly eusocial, since their social behavior depends largely on their resources and environment. Some mammals in the Carnivora and Primates have eusocial tendencies, especially meerkats ( Suricata suricatta ) and dwarf mongooses ( Helogale parvula ). These show cooperative breeding and marked reproductive skews. In
8190-399: The individuals in the colony are to some degree specialized for tasks, a division of labor . Eusociality is a rare but widespread phenomenon in species in at least seven orders in the animal kingdom , as shown in the phylogenetic tree (non-eusocial groups not shown). All species of termites are eusocial, and it is believed that they were the first eusocial animals to evolve, sometime in
8295-406: The large size of their colonies, and surplus production and storage of honey , distinguishing their hives as a prized foraging target of many animals, including honey badgers , bears and human hunter-gatherers . Only 8 surviving species of honey bee are recognized, with a total of 43 subspecies , though historically 7 to 11 species are recognized. Honey bees represent only a small fraction of
8400-417: The largest group of eusocial insects, including ants , bees , and wasps —divided into castes: reproductive queens , drones , more or less sterile workers, and sometimes also soldiers that perform specialized tasks. In the well-studied social wasp Polistes versicolor , dominant females perform tasks such as building new cells and ovipositing, while subordinate females tend to perform tasks like feeding
8505-561: The larvae and foraging. The task differentiation between castes can be seen in the fact that subordinates complete 81.4% of the total foraging activity, while dominants only complete 18.6% of the total foraging. Eusocial species with a sterile caste are sometimes called hypersocial. While only a moderate percentage of species in bees (families Apidae and Halictidae ) and wasps ( Crabronidae and Vespidae ) are eusocial, nearly all species of ants ( Formicidae ) are eusocial. Some major lineages of wasps are mostly or entirely eusocial, including
8610-561: The loss of their queen. The levels of two of the aliphatic compounds increase rapidly in virgin queens within the first week after emergence from the pupa , consistent with their roles as sex attractants during the mating flight. Once a queen is mated and begins laying eggs, she starts producing the full blend of compounds. In several ant species, reproductive activity is associated with pheromone production by queens. Mated egg-laying queens are attractive to workers, whereas young winged virgin queens elicit little or no response. Among ants,
8715-610: The majority of social bee diversity, have their lowest diversity in the tropics, being largely limited to temperate regions. Once pre-adaptations such as group formation, nest building, high cost of dispersal, and morphological variation are present, between-group competition has been suggested as a driver of the transition to advanced eusociality. M. A. Nowak, C. E. Tarnita, and E. O. Wilson proposed in 2010 that since eusociality produces an extremely altruistic society, eusocial groups should out-reproduce their less cooperative competitors, eventually eliminating all non-eusocial groups from
8820-591: The mite population may increase to such an extent that the hive will succumb to the diseases and deformities caused by the mites. It was widely believed that the mites drank the blood of bees. However, a 2018 study Article in PNAS: "Linking pesticides and gut health in bees" showed that they actually feed on the fat body tissue of live bees, not the blood. Mite treatment is accomplished by several methods, including treatment strips and acid vaporization. Eusocial Eusociality ( Greek εὖ eu "good" and social )
8925-423: The mite, temperature treatments are possible, but also distorts wax of the honeycombs. Chemical fumigants, particularly CO 2 , are also used. Varroa mites are arguably the biggest threat to honey bees in the United States. These mites invade hives and reproduce by laying eggs on pupa. The hatching mites eat away at the pupa, causing deformities as well as spreading disease. If not detected and treated early on,
9030-532: The most common circulating sugar in hemolymph is trehalose which is a disaccharide consisting of two glucose molecules. Adult worker honey bees require 4 mg of utilizable sugars per day and larvae require about 59.4 mg of carbohydrates for proper development. Honey bees require water to maintain osmotic homeostasis, prepare liquid brood food, and to cool the hive through evaporation. A colony's water needs can generally be met by nectar foraging as it has high water content. Occasionally on hot days or when nectar
9135-465: The number of brood reared. Honey bees ingest phytosterols from pollen to produce 24-methylenecholesterol and other sterols as they cannot directly synthesize cholesterol from phytosterols. Nurse bees have the ability to selectively transfer sterols to larvae through brood food. Nectar is collected by foraging worker bees as a source of water and carbohydrates in the form of sucrose. The dominant monosaccharides in honey bee diets are fructose and glucose but
9240-501: The only extant members of the tribe Apini. Today's honey bees constitute three clades : Micrapis (the dwarf honey bees), Megapis (the giant honey bee), and Apis (the western honey bee and its close relatives). Most species have historically been cultured or at least exploited for honey and beeswax by humans indigenous to their native ranges. Only two species have been truly domesticated : Apis mellifera and Apis cerana . A. mellifera has been cultivated at least since
9345-401: The opportunity. One plant, the epiphytic staghorn fern, Platycerium bifurcatum ( Polypodiaceae ), may exhibit a primitive form of eusocial behavior amongst clones. The evidence for this is that individuals live in colonies, where they are structured in different ways, with fronds of differing size and shape, to collect and store water and nutrients for the colony to use. At the top of
9450-617: The other lineages, but do not seem to have diverged from each other a long time before the Neogene . Apis florea have smaller wing spans than its sister species. Apis florea are also completely yellow with the exception of the scutellum of workers, which is black. Two species are recognized in the subgenus Megapis . They usually build single or a few exposed combs on high tree limbs, on cliffs, and sometimes on buildings. They can be very fierce. Periodically robbed of their honey by human "honey hunters", colonies are easily capable of stinging
9555-495: The place of origin of the genus, only that the bees were present in Europe by that time. Few fossil deposits are known from South Asia, the suspected region of honey bee origin, and fewer still have been thoroughly studied. No Apis species existed in the New World during human times before the introduction of A. mellifera by Europeans. Only one fossil species is documented from the New World, Apis nearctica , known from
9660-627: The queen pheromone system of the fire ant Solenopsis invicta includes both releaser and primer pheromones. A queen recognition (releaser) pheromone is stored in the poison sac along with three other compounds. These compounds elicit a behavioral response from workers. Several primer effects have also been demonstrated. Pheromones initiate reproductive development in new winged females, called female sexuals. These chemicals inhibit workers from rearing male and female sexuals, suppress egg production in other queens of multiple queen colonies, and cause workers to execute excess queens. These pheromones maintain
9765-626: The reproductive output of other individuals that share their genes, especially their close relatives. Natural selection favors individuals to help their relatives when the cost of helping is less than the benefit gained by their relative multiplied by the fraction of genes that they share, i.e. when Cost < relatedness * Benefit . W. D. Hamilton suggested in 1964 that eusociality could evolve more easily among haplodiploid species such as Hymenoptera, because of their unusual relatedness structure. In haplodiploid species, females develop from fertilized eggs and males develop from unfertilized eggs. Because
9870-457: The result of allopatric speciation , their distribution later converging. Given that A. florea is more widely distributed and A. andreniformis is considerably more aggressive, honey is, if at all, usually harvested from the former only. They are the most ancient extant lineage of honey bees, maybe diverging in the Bartonian (some 40 million years ago or slightly later) from
9975-535: The roles of the sole reproducers, while soldiers and workers work together to create and maintain a living situation favorable for the brood. Queens produce multiple queen pheromones to create and maintain the eusocial state in their colonies; they may also eat eggs laid by other females or exert dominance by fighting. There are two eusocial rodents : the naked mole-rat and the Damaraland mole-rat . Some shrimps , such as Synalpheus regalis , are eusocial. E. O. Wilson and others have claimed that humans have evolved
10080-479: The roughly 20,000 known species of bees. The best known honey bee is the western honey bee , ( Apis mellifera ), which was domesticated for honey production and crop pollination . The only other domesticated bee is the eastern honey bee ( Apis cerana ), which occurs in South , Southeast , and East Asia . Only members of the genus Apis are true honey bees, but some other types of bees produce and store honey and have been kept by humans for that purpose, including
10185-574: The solitary state. Pheromones play an important role in the physiological mechanisms of eusociality. Enzymes involved in the production and perception of pheromones were important for the emergence of eusociality within both termites and hymenopterans. The best-studied queen pheromone system in social insects is that of the honey bee Apis mellifera . Queen mandibular glands produce a mixture of five compounds, three aliphatic and two aromatic , which control workers. Mandibular gland extracts inhibit workers from constructing queen cells, which can delay
10290-402: The subfamilies Polistinae and Vespinae . The corbiculate bees (subfamily Apinae of family Apidae ) contain four tribes of varying degrees of sociality: the highly eusocial Apini (honey bees) and Meliponini (stingless bees), primitively eusocial Bombini (bumble bees), and the mostly solitary or weakly social Euglossini (orchid bees). Eusociality in these families is sometimes managed by
10395-566: The time of the building of the Egyptian pyramids , and only that species has been moved extensively beyond its native range. Apis florea and Apis andreniformis are small honey bees of southern and southeastern Asia. They make very small, exposed nests in trees and shrubs. Their stings are often incapable of penetrating human skin, so the hive and swarms can be handled with minimal protection. They occur largely sympatrically , though they are very distinct evolutionarily and are probably
10500-749: The upper Jurassic period (~150 million years ago). The other orders shown contain both eusocial and non-eusocial species, including many lineages where eusociality is inferred to be the ancestral state. Thus the number of independent evolutions of eusociality ( clades ) is not known. The major eusocial groups are shown in boldface in the phylogenetic tree. Mole-rats [REDACTED] Synalpheus spp. [REDACTED] [REDACTED] Kladothrips spp. [REDACTED] various Aphids [REDACTED] Austroplatypus incompertus [REDACTED] many Vespidae (wasps) [REDACTED] [REDACTED] many Bees [REDACTED] Staghorn fern Platycerium bifurcatum [REDACTED] Prior to
10605-490: The western honey bee and deliberately transferred them on a global scale, diseases simultaneously spread and harmed managed colonies. Colony losses have occurred periodically throughout history. Fungus, mites, and starvation have all been thought to be the cause of the deaths. Limited occurrences resembling CCD were documented as early as 1869. Colony collapses were called "May Disease" in Colorado in 1891 and 1896. Starting in
10710-411: The winter is acceptable and more than three-fifths of beekeepers surveyed said their losses were higher than that in 2022-2023. Acarapis woodi (or "tracheal mites") are parasitic mites which live and reproduce in adult bees' tracheae, or respiratory tubes, piercing the tube walls with their mouthparts to feed on haemolymph. To infest new hosts, the mites must find newly emerged bees; after three days,
10815-419: The workers perform tasks within the nest such as provisioning cell broods, colony cleaning, and nectar reception and dehydration. Once older than 40 days, S. postica workers move outside the nest for colony defense and foraging. In Lasioglossum aeneiventre , a halictid bee from Central America, nests may be headed by more than one female; such nests have more cells, and the number of active cells per female
10920-422: Was followed by at least one reversal to solitarity, giving a total of at least nine reversals. In a few species, solitary and eusocial colonies appear simultaneously in the same population, and different populations of the same species may be fully solitary or eusocial. This suggests that eusociality is costly to maintain, and can only persist when ecological variables favor it. Disadvantages of eusociality include
11025-511: Was then discovered in a group of chordates , the mole-rats. Further research distinguished another possibly important criterion for eusociality, "the point of no return". This is characterized by having individuals fixed into one behavioral group, usually before reproductive maturity. This prevents them from transitioning between behavioral groups, and creates a society with individuals truly dependent on each other for survival and reproductive success. For many insects, this irreversibility has changed
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