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Honey bee
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For other uses, see Honey bee (disambiguation).
Honey bees
Temporal range: Oligocene–Recent
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Western honey bee carrying pollen back to the hive
Scientific classification
Kingdom: Animalia
Clade: Euarthropoda
Class: Insecta
Order: Hymenoptera
Family: Apidae
Subfamily: Apinae
Tribe: Apini
Latreille, 1802
Genus: Apis
Linnaeus, 1758
Species
†Apis lithohermaea
†Apis nearctica
Subgenus Micrapis:
Apis andreniformis
Apis florea
Subgenus Megapis:
Apis dorsata
Subgenus Apis:
Apis cerana
Apis koschevnikovi
Apis mellifera
Apis nigrocincta
A honey bee (or honeybee) is any member of the genus Apis, primarily distinguished by the production and storage of honey and the construction of perennial, colonial nests from wax. In the early 21st century, only seven species of honey bee are recognized, with a total of 44 subspecies,[1] though historically six to eleven species are recognized. The best known honey bee is the Western honey bee which has been domesticated for honey production and crop pollination. Honey bees represent only a small fraction of the roughly 20,000 known species of bees.[2] Some other types of related bees produce and store honey, including the stingless honey bees, but only members of the genus Apis are true honey bees. The study of bees, which includes the study of honey bees, is known as melittology.
Contents
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1 Etymology and name
2 Origin, systematics and distribution
2.1 Genetics
2.2 Micrapis
2.3 Megapis
2.4 Apis
2.5 Borneo honey bee
2.6 Eastern honey bee
2.7 Western honey bee
2.8 Africanized bee
3 Living and fossil honey bees (Apini: Apis)
4 Life cycle
4.1 Life cycle
4.2 Winter survival
5 Pollination
6 Nutrition
7 Parasites
7.1 Galleria mellonella
7.1.1 Management
8 Beekeeping
8.1 Colony collapse disorder
9 Bee products
9.1 Honey
9.2 Beeswax
9.3 Pollen
9.4 Bee bread
9.5 Propolis
9.6 Royal jelly
10 Sexes and castes
10.1 Drones
10.2 Workers
10.3 Queens
11 Defense
12 Competition
13 Communication
14 Symbolism
15 Gallery
16 See also
17 Notes
18 References
19 Further reading
20 External links
Etymology and name[edit]
The genus name Apis is Latin for "bee".[3]
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, as it is a kind or type of bee, whereas it is incorrect to run the two words together as in dragonfly or butterfly, because the latter are not flies.[4] 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.[5][6][7] Nonetheless, compounds gradually solidify in the orthography of natural languages in ways that do not always comply with prescription.
Origin, systematics and distribution[edit]
Distribution of honey bees around the world
Morphology of a female honey bee
Honey bees appear to have their center of origin in South and Southeast Asia (including 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.[8]
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 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 a single 14 million-year-old specimen from Nevada.[9]
The close relatives of modern honey bees – e.g. bumblebees and stingless bees – are also social to some degree, and social behavior seems a plesiomorphic 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.
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 indica. A. mellifera has been cultivated at least since the time of the building of the Egyptian pyramids, and only that species has been moved extensively beyond its native range.
Honey bees are the only extant members of the tribe Apini. Today's honey bees constitute three clades: Micrapis (dwarf honey bees), Megapis (giant honey bee), and Apis (domestic honey bees and close relatives).[1][10]
Genetics[edit]
Main article: Apis mellifera
The genome of Apis has been mapped.
The chromosome counts of female bees for the three clades are:
Clade 2 N
Micrapis 16
Megapis 16
Apis 32
Drones (males) are produced from unfertilized eggs, so they represent only the DNA of the queen that laid the eggs, i.e. have only a mother. Drones of all bee genera have 1 N chromosome counts – half of those shown above.
Workers and queens (both female) result from fertilized eggs, so have both a mother and a father. Arrhenotokous parthenogenesis, a modified form of parthenogenesis, controls sex differentiation. The sex allele is polymorphic, and so long as two different variants are present, a female bee results. If both sex alleles are identical, diploid drones are produced. Honey bees detect and destroy diploid drones after the eggs hatch.
Queens typically mate with multiple drones on more than one mating flight. Once mated, they lay eggs and fertilize them as needed from sperm stored in the spermatheca. Since the number of sex alleles is limited at a regional level, a queen will most likely mate with one or more drones having sex alleles identical with one of the sex alleles in the queen. The queen, then, typically produces a percentage of diploid drone eggs.[a]
Micrapis[edit]
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 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 the other lineages, but do not seem to have diverged from each other a long time before the Neogene.[10] Apis florea have smaller wing spans than its sister species.[12] Apis florea are also completely yellow with the exception of the scutellum of workers, which is black.[12]
Megapis[edit]
One species is recognized in the subgenus Megapis. It usually builds 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 a human being to death if provoked.
Apis dorsata, the giant honey bee, is native and widespread across most of South and Southeast Asia.
A. d. binghami, the Indonesian giant honey bee, is classified as the Indonesian subspecies of the giant honey bee or a distinct species; in the latter case, A. d. breviligula and / or other lineages would probably also have to be considered species.[13]
A. d. laboriosa, the Himalayan giant honey bee, was initially described as a distinct species. Later, it was included in A. dorsata as a subspecies[1] based on the biological species concept, though authors applying a genetic species concept have suggested it should be considered a separate species.[10] Essentially restricted to the Himalayas, it differs little from the giant honey bee in appearance, but has extensive behavioral adaptations that enable it to nest in the open at high altitudes despite low ambient temperatures. It is the largest living honey bee.
Apis[edit]
Eastern Apis species include three or four species, including A. koschevnikovi and A. cerana. The genetics of the western honey bee A. mellifera are unclear. The domesticated species are A. cerana indica and A. mellifera
Borneo honey bee[edit]
The reddish Koschevnikov's bee (Apis koschevnikovi) from Borneo is clearly a distinct species; it probably derives from the first colonization of the island by cave-nesting honey bees.
Eastern honey bee[edit]
Apis cerana, the eastern honey bee proper, is the traditional honey bee of southern and eastern Asia. It was domesticated as subspecies A. c. indica 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 A. c. nuluensis and Apis nigrocincta from the Philippines to satisfaction; the most recent hypothesis is that these are indeed distinct species, but that A. cerana is still paraphyletic, consisting of several separate species.[10]
Western honey bee[edit]
The European honey bee may have originated from eastern Africa. This bee is pictured in Tanzania.
A. mellifera, the most common domesticated[14] species, was the third insect to have its genome mapped. It seems to have originated in eastern tropical Africa and spread from there to Northern Europe and eastwards into Asia to the Tien 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, hybrid strains, such as the Buckfast bee, have been bred. Behavior, color, 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 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 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 the exact relationships of many A. mellifera subspecies.[10]
Apis mellifera is not native to the Americas, so was not present upon the arrival of the European explorers and colonists. However, other native bee species were kept and traded by indigenous peoples. In 1622, European colonists brought the dark bee (A. m. mellifera) to the Americas, followed later by Italian bees (A. m. ligustica) and others. Many of the crops that depend on honey bees for pollination have also been imported since colonial times. Escaped swarms (known as “wild” 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 the early 1850s.[15]
An African bee in Tanzania
Africanized bee[edit]
Main article: Africanized bee
Africanized bees (known colloquially as "killer bees") are hybrids between European stock and one of the African subspecies A. m. scutellata; they are often more aggressive than European bees and do not create as much of a honey surplus, but are more resistant to disease and are better foragers.[citation needed] Originating by accident in Brazil, they have spread to North America and constitute a pest in some regions. However, these strains do not overwinter well, so are not often found in the colder, more northern parts of North America. The original breeding experiment for which the African bees were brought to Brazil in the first place has continued (though not as intended). Novel hybrid strains of domestic and redomesticated Africanized bees combine high resilience to tropical conditions and good yields. They are popular among beekeepers in Brazil.
Living and fossil honey bees (Apini: Apis)[edit]
Tribe Apini Latreille[16]
Genus Apis Linnaeus (s. lato)
henshawi species group (†Priorapis Engel, †Synapis Cockerell)
†A. vetusta Engel
†A. henshawi Cockerell
†A. petrefacta (Říha)
†A. miocenica Hong
†A. “longtibia” Zhang
†A. “Miocene 1”
armbrusteri species group (†Cascapis Engel)
†A. armbrusteri Zeuner
†A. nearctica, sp. Nov.
florea species group (Micrapis Ashmead)
A. florea Fabricius
A. andreniformis Smith
dorsata species group (Megapis Ashmead)
†A. lithohermaea Engel
A. dorsata Fabricius
mellifera species group (Apis Linnaeus s. stricto)
mellifera subgroup
A. mellifera Linnaeus (Apis Linnaeus s. strictissimo)
cerana subgroup (Sigmatapis Maa)
A. cerana Fabricius
A. nigrocincta Smith
A. koschevnikovi Enderlein
Life cycle[edit]
As in a few other types of eusocial bees, a colony generally contains one queen bee, a fertile female; seasonally up to a few thousand drone bees, or fertile males;[17] and tens of thousands of sterile female worker bees. Details vary among the different species of honey bees, but common features include:
Eggs are laid singly in a cell in a wax honeycomb, produced and shaped by the worker bees. Using her spermatheca, the queen can choose to fertilize the egg she is laying, usually depending on into which cell she is laying. Drones develop from unfertilised eggs and are haploid, while females (queens and worker bees) develop from fertilised eggs and are diploid. Larvae are initially fed with royal jelly produced by worker bees, later switching to honey and pollen. The exception is a larva fed solely on royal jelly, which will develop into a queen bee. The larva undergoes several moultings before spinning a cocoon within the cell, and pupating.
Young worker bees, sometimes called "nurse bees", clean the hive and feed the larvae. When their royal jelly-producing glands begin to atrophy, they begin building comb cells. They progress to other within-colony tasks as they become older, such as receiving nectar and pollen from foragers, and guarding the hive. Later still, a worker takes her first orientation flights and finally leaves the hive and typically spends the remainder of her life as a forager.
Worker bees cooperate to find food and use a pattern of "dancing" (known as the bee dance or waggle dance) to communicate information regarding resources with each other; this dance varies from species to species, but all living species of Apis exhibit some form of the behavior. If the resources are very close to the hive, they may also exhibit a less specific dance commonly known as the "round dance".
Honey bees also perform tremble dances, which recruit receiver bees to collect nectar from returning foragers.
Virgin queens go on mating flights away from their home colony to a drone congregation area, and mate with multiple drones before returning. The drones die in the act of mating. Queen honey bees do not mate with drones from their home colony.
Colonies are established not by solitary queens, as in most bees, but by groups known as "swarms", which consist of a mated queen and a large contingent of worker bees. This group moves en masse to a nest site which was scouted by worker bees beforehand and whose location is communicated with a special type of dance. Once the swarm arrives, they immediately construct a new wax comb and begin to raise new worker brood. This type of nest founding is not seen in any other living bee genus, though several groups of vespid wasps also found new nests by swarming (sometimes including multiple queens). Also, stingless bees will start new nests with large numbers of worker bees, but the nest is constructed before a queen is escorted to the site, and this worker force is not a true "swarm".
Life cycle[edit]
A coloured dot applied by a beekeeper identifies the queen
Honey bee eggs shown in opened wax cells
Eggs and larvae
Drone pupae
Emergence of a black bee (A. m. mellifera)
Winter survival[edit]
In cold climates, honey bees stop flying when the temperature drops below about 10 °C (50 °F) and crowd into 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 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).[18] In addition, certain bees, including 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.[19]
Pollination[edit]
Main articles: Pollination management and List of crop plants pollinated by bees
Species of Apis are generalist floral visitors, and pollinate a large variety of plants, but by no means all plants. Of all the honey bee species, only A. mellifera has been used extensively for commercial pollination of crops and other plants. The value of these pollination services is commonly measured in the billions of dollars. Bees collect 66 pounds (30 kg) of pollen per year per hive.[20]
Nutrition[edit]
Honey bees obtain all of their nutritional requirements from 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.[21] The rearing of one larva requires 125-187.5 mg pollen or 25-37.5 mg protein for proper development.[21] 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.[22] 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 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.[22]
A forager collecting pollen
Pollen is also a lipid source for honey bees ranging from 0.8% to 18.9%.[21] 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 the number of brood reared.[21] 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.[21]
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 the most common circulating sugar in hemolymph is trehalose which is a disaccharide consisting of two glucose molecules.[23] Adult worker honey bees require 4 mg of utilizable sugars per day and larvae require about 59.4 mg of carbohydrates for proper development.[21]
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 is limited, foragers will collect water from streams or ponds to meet the needs of the hive.[24]
Parasites[edit]
Galleria mellonella[edit]
Larval stages of G. mellonella parasitize both wild and cultivated honeybees. Eggs are laid within the hive, and the larva that hatch tunnel through and destroy the honeycombs that contain honeybee 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. Finally, both G. mellonella adults and larvae can be vectors for pathogens that can infect honeybees, including the Israeli acute paralysis virus (IAPV) and the black queen cell virus (BQCV).[25]
Management[edit]
Temperature treatments are possible, but also distorts wax of the honeycombs. Chemical fumigants, particularly CO2,are also used. [25]
Beekeeping[edit]
Main article: Beekeeping
Two species of honey bee, A. mellifera and A. cerana indica, are often maintained, fed, and transported by beekeepers. Modern hives also enable beekeepers to transport bees, moving from field to field as 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.
Colony collapse disorder[edit]
Main article: Colony collapse disorder
Since 2007, abnormally high die-offs (30–70% of hives) of European honey bee colonies have occurred in North America. This has been dubbed "colony collapse disorder" (CCD) and was at first unexplained.[26] It seems to be caused by a combination of factors rather than a single pathogen or poison, possibly including neonicotinoid pesticides[27] or Israeli acute paralysis virus.[28]
Bee products[edit]
Honey[edit]
Main article: Honey
Honey is the complex substance made when bees ingest nectar, process it, and store the substance into honey combs.[29] All living species of Apis have had their honey gathered by indigenous peoples for consumption. A. mellifera and A. cerana are the only species that have had their honey harvested for commercial purposes.
Beeswax[edit]
Main article: Beeswax
Worker bees of a certain age secrete beeswax from a series of glands on their abdomens.[30] They use the wax to form the walls and caps of the comb. As with honey, beeswax is gathered by humans for various purposes.
Pollen[edit]
Main article: Bee pollen
Bees collect pollen in their pollen baskets and carry it back to the hive.[31] In the hive, pollen is used as a protein source necessary during brood-rearing. In certain environments, excess pollen can be collected from the hives of A. mellifera and A. cerana. It is often eaten as a health supplement. It also has been used with moderate success as a source of pollen for hand pollination.
Bee bread[edit]
Worker bees combine pollen, honey and glandular secretions and allow it to ferment in the comb to make bee bread. The fermentation process releases additional nutrients from the pollen and can produce antibiotics and fatty acids which inhibit spoilage. Bee bread is eaten by nurse bees (younger workers) who then produce the protein-rich royal jelly needed by the queen and developing larvae in their hypopharyngeal glands.
Propolis[edit]
Main article: Propolis
Propolis is a resinous mixture collected by honey bees from tree buds, sap flows or other botanical sources, which is used as a sealant for unwanted open spaces in the hive.[32] Although propolis is alleged to have health benefits (tincture of Propolis is marketed as a cold and flu remedy), it may cause severe allergic reactions in some individuals.[33] Propolis is also used in wood finishes, and gives a Stradivarius violin its unique red color.[34]
Royal jelly[edit]
Main article: Royal jelly
Royal jelly is a honey-bee secretion used to nourish the larvae.[35] It is marketed for its alleged but unsupported claims of health benefits.[36][37] On the other hand, it may cause severe allergic reactions in some individuals.[38]
Sexes and castes[edit]
Sexes and roles in a colony of honey bees
Honey bees have three castes: drones, workers, and queens.[39][40] Drones are male, while workers and queens are female.[40]
Drones[edit]
Honey bees have a haplodiploid system of sex determination.
Main article: Drone (bee)
Males, or drones, are typically haploid, having only one set of chromosomes, and primarily exist for the purpose of reproduction.[40] They are produced by the queen if she chooses not to fertilize an egg or by an unfertilized laying worker. Diploid drones may be produced if an egg is fertilized but is homozygous for the sex-determination allele. Drones take 24 days to develop, and may be produced from summer through to autumn, numbering as many as 500 per hive.[40] They are expelled from the hive during the winter months when the hive's primary focus is warmth and food conservation.[40] Drones have large eyes used to locate queens during mating flights. They do not defend the hive or kill intruders, and do not have a stinger.[41]
Workers[edit]
Main article: Worker bee
Workers have two sets of chromosomes.[42] They are produced from an egg that the queen has selectively fertilized from stored sperm. Workers typically develop in 21 days. A typical colony may contain as many as 60,000 worker bees.[40] Workers exhibit a wider range of behaviors than either queens or drones. Their duties change upon the age of the bee in the following order (beginning with cleaning out their own cell after eating through their capped brood cell): feed brood, receive nectar, clean hive, guard duty, and foraging.[40][41] Some workers engage in other specialized behaviors, such as "undertaking" (removing corpses of their nestmates from inside the hive).[41]
Workers have morphological specializations, including the pollen basket (corbicula),[43] abdominal glands that produce beeswax, brood-feeding glands, and barbs on the sting. Under certain conditions (for example, if the colony becomes queenless), a worker may develop ovaries.
Queens[edit]
Main article: Queen bee
Queen honey bees are created when worker bees feed a single female larvae an exclusive diet of a food called "royal jelly".[40][41] Queens are produced in oversized cells and develop in only 16 days; they differ in physiology, morphology, and behavior from worker bees. In addition to the greater size of the queen, she has a functional set of ovaries, and a spermatheca, which stores and maintains sperm after she has mated. Apis queens practice polyandry, with one female mating with multiple males. The highest documented mating frequency for an Apis queen is in Apis nigrocincta, where queens mate with an extremely high number of males with observed numbers of different matings ranging from 42 to 69 drones per queen.[44] The sting of queens is not barbed like a worker's sting, and queens lack the glands that produce beeswax. Once mated, queens may lay up to 2,000 eggs per day.[41] They produce a variety of pheromones that regulate behavior of workers, and helps swarms track the queen's location during the swarming.[41]
Defense[edit]
Main article: Bee sting
Apis cerana japonica forming a ball around two hornets: The body heat trapped by the ball will overheat and kill the hornets.
All honey bees live in colonies where the workers sting intruders as a form of defense, and alarmed bees release a pheromone that stimulates the attack response in other bees. The different species of honey bees are distinguished from all other bee species (and virtually all other Hymenoptera) by the possession of small barbs on the sting, but these barbs are found only in the worker bees. The sting and associated venom sac of honey bees are also modified so as to pull free of the body once lodged (autotomy), and the sting apparatus has its own musculature and ganglion, which allows it to keep delivering venom once detached. The gland which produces the alarm pheromone is also associated with the sting apparatus. The embedded stinger continues to emit additional alarm pheromone after it has torn loose; other defensive workers are thereby attracted to the sting site. The worker dies after the sting becomes lodged and is subsequently torn loose from the bee's abdomen. The honey bee's venom, known as apitoxin, carries several active components, the most abundant of which is melittin, and the most destructive is phospholipase A2.
This complex apparatus, including the barbs on the sting, is thought to have evolved specifically in response to predation by vertebrates, as the barbs do not usually function (and the sting apparatus does not detach) unless the sting is embedded in fleshy tissue. While the sting can also penetrate the membranes between joints in the exoskeleton of other insects (and is used in fights between queens), in the case of Apis cerana japonica, defense against larger insects such as predatory wasps (e.g. Asian giant hornet) is usually performed by surrounding the intruder with a mass of defending worker bees, which vibrate their muscles vigorously to raise the temperature of the intruder to a lethal level ("balling").[45] Previously, heat alone was thought to be responsible for killing intruding wasps, but recent experiments have demonstrated the increased temperature in combination with increased carbon dioxide levels within the ball produce the lethal effect.[46][47] This phenomenon is also used to kill a queen perceived as intruding or defective, an action known to beekeepers as 'balling the queen', named for the ball of bees formed.
Defense can vary based on the habitat of the bee. In the case of those honey bee species with open combs (e.g., A. dorsata), would-be predators are given a warning signal that takes the form of a "Mexican wave" that spreads as a ripple across a layer of bees densely packed on the surface of the comb when a threat is perceived, and consists of bees momentarily arching their bodies and flicking their wings.[48] In cavity dwelling species such as Apis cerana, Apis mellifera, and Apis nigrocincta, entrances to these cavities are guarded and checked for intruders in incoming traffic. Another act of defense against nest invaders, particularly wasps, is "body shaking," a violent and pendulum like swaying of the abdomen, performed by worker bees.[49]
Competition[edit]
With an increased number of honey bees in a specific area due to beekeeping, domesticated bees and native wild bees often have to compete for the limited habitat and food sources available.[50] European bees may become defensive in response to the seasonal arrival of competition from other colonies, particularly Africanized bees which may be on the offence and defense year round due to their tropical origin.[51] In the United Kingdom, honey bees are known to compete with Bombus hortorum, a bumblebee species, because they forage at the same sites. To resolve the issue and maximize both their total consumption during foraging, bumblebees forage early in the morning, while honey bees forage during the afternoon.[52]
Communication[edit]
Main article: Bee learning and communication
Honey bees are known to communicate through many different chemicals and odors, as is common in insects. They also rely on a sophisticated dance language that conveys information about the distance and direction to a specific location (typically a nutritional source, e.g., flowers or water). The dance language is also used during the process of reproductive fission, or swarming, when scouts communicate the location and quality of nesting sites.[53]
The details of the signalling being used vary from species to species; for example, the two smallest species, Apis andreniformis and A. florea, dance on the upper surface of the comb, which is horizontal (not vertical, as in other species), and worker bees orient the dance in the actual compass direction of the resource to which they are recruiting.
Apis mellifera carnica honey bees use their antennae asymmetrically for social interactions with a strong lateral preference to use their right antennae.[54][55]
There has been speculation as to honeybee consciousness.[56]
Symbolism[edit]
Main article: Bee (mythology)
The bee was revived as a symbol of government by Emperor Napoleon I of France.[57] Both the Hindu Atharva Veda[58] and the ancient Greeks associated lips anointed with honey with the gift of eloquence and even of prescience. The priestess at Delphi was the "Delphic Bee". The Quran has a chapter titled "The Bee". A community of honey bees has often been employed by political theorists as a model of human society, from Aristotle and Plato to Virgil[59] and Seneca; in Erasmus and Shakespeare; in Marx and Tolstoy.[60] Honey bees, signifying immortality and resurrection, were royal emblems of the Merovingians. The bee also is the heraldic emblem of the Barberini.[citation needed] The state of Utah is called the "Beehive State", the state emblem is the beehive, the state insect is the honey bee, and a beehive and the word "industry" appear on both the state flag and seal.[61][62][63][64]
Gallery[edit]
Foragers loaded with pollen on the hive landing board
Eastern honey bee (A. cerana) in Hong Kong
Giant honey bee (A. dorsata)
Honey bee visiting flowers
A colony of giant honey bees on their comb
Play media
Pollinating flowers
Genus Apis
See also[edit]
Bees and toxic chemicals
Honey bee life cycle
More Than Honey – a 2012 Swiss documentary film about honey bees
Honeybee Starvation
Notes[edit]
Jump up ^ Work by Lechner in 2013 showed approximately 50 sex alleles in a population of Apis mellifera and estimated worldwide total number of sex alleles in A. mellifera between 116 and 145. There are likely more sex alleles in Apis cerana given wider distribution and larger number of geographical races. Honeybees are highly sensitive to inbreeding in part because duplication of the sex allele causes significant numbers of diploid drone eggs to be laid and subsequently destroyed as they hatch.[11]
References[edit]
^ Jump up to: a b c Michael S. Engel (1999). "The taxonomy of recent and fossil honey bees (Hymenoptera: Apidae: Apis)". Journal of Hymenoptera Research. 8: 165–196.
Jump up ^ "Bees - Facts About Bees - Types of Bees - PestWorldforKids.org". pestworldforkids.org. Retrieved 2016-04-26.
Jump up ^ Douglas Harper (2006). "Online Etymology Dictionary". Retrieved 2016-02-27.
Jump up ^ Robert E. Snodgrass (1984). Anatomy of the Honey Bee. Cornell University Press. p. vii. ISBN 0-8014-9302-1.
Jump up ^ "ITIS". Retrieved February 26, 2016.
Jump up ^ "Entomological Society of America Common Names of Insects Database". Retrieved February 21, 2016.
Jump up ^ "Tree of Life Web Project". Retrieved 2016-02-25.
Jump up ^ Deborah R. Smith; Lynn Villafuerte; Gard Otisc; Michael R. Palmer (2000). "Biogeography of Apis cerana F. and A. nigrocincta Smith: insights from mtDNA studies" (PDF). Apidologie. 31 (2): 265–279. doi:10.1051/apido:2000121. Archived from the original (PDF) on February 29, 2012.
Jump up ^ Michael S. Engel; I. A. Hinojosa-Diaz; A. P. Rasnitsyn (2009). "A honey bee from the Miocene of Nevada and the biogeography of Apis (Hymenoptera: Apidae: Apini)". Proceedings of the California Academy of Sciences. 60 (3): 23–38.
^ Jump up to: a b c d e Maria C. Arias; Walter S. Sheppard (2005). "Phylogenetic relationships of honey bees (Hymenoptera:Apinae:Apini) inferred from nuclear and mitochondrial DNA sequence data". Molecular Phylogenetics and Evolution. 37 (1): 25–35. doi:10.1016/j.ympev.2005.02.017. PMID 16182149.
Maria C. Arias; Walter S. Sheppard (2005). "Corrigendum to "Phylogenetic relationships of honey bees (Hymenoptera:Apinae:Apini) inferred from nuclear and mitochondrial DNA sequence data"". Molecular Phylogenetics and Evolution. 40 (1): 315. doi:10.1016/j.ympev.2006.02.002.
Jump up ^ Lechner, Sarah; Ferretti, Luca; Schöning, Caspar; Kinuthia, Wanja; Willemsen, David; Hasselmann, Martin (2014). "Nucleotide Variability at Its Limit? Insights into the Number and Evolutionary Dynamics of the Sex-Determining Specificities of the Honey Bee Apis mellifera". Molecular Biology and Evolution. 31 (2): 272–287. doi:10.1093/molbev/mst207.
^ Jump up to: a b Wongsiri, S., et al. "Comparative biology of Apis andreniformis and Apis florea in Thailand." Bee World 78.1 (1997): 23-35.
Jump up ^ Nathan Lo; Rosalyn S. Gloag; Denis L. Anderson; Benjamin P. Oldroyd (2009). "A molecular phylogeny of the genus Apis suggests that the Giant Honey Bee of the Philippines, A. breviligula Maa, and the Plains Honey Bee of southern India, A. indica Fabricius, are valid species". Systematic Entomology. 35 (2): 226–233. doi:10.1111/j.1365-3113.2009.00504.x.
Jump up ^ mostly domesticated or more accurately, "managed"
Jump up ^ Head RJ (2008). "A Brief Survey of Ancient Near Eastern Beekeeping; A Final Note". The FARMS Review.
Jump up ^ Michael S. Engel, Ismael A. Hinojosa-Díaz & Alexandr P. Rasnitsyn (2009). "A honey bee from the Miocene of Nevada and the biogeography of Apis (Hymenoptera: Apidae: Apini)" (PDF). Proceedings of the California Academy of Sciences. 4. 60 (3): 23–38.
Jump up ^ James L. Gould; Carol Grant Gould (1995). The Honey Bee. Scientific American Library. p. 19. ISBN 978-0-7167-6010-8.
Jump up ^ "What do bees do in the winter?". Retrieved 12 March 2016.
Jump up ^ Oldroyd, Benjamin P.; Wongsiri, Siriwat (2006). Asian Honey Bees (Biology, Conservation, and Human Interactions). Cambridge, Massachusetts and London, England: Harvard University Press.ISBN 0674021940.
Jump up ^ "Facts about Honeybees". Back Yard Beekeepers Association. 2017.
^ Jump up to: a b c d e f Brodschneider, Robert; Crailsheim, Karl (2010-05-01). "Nutrition and health in honey bees". Apidologie. 41 (3): 278–294. doi:10.1051/apido/2010012. ISSN 0044-8435.
^ Jump up to: a b Anderson, Leroy M; Dietz, A. (1976). "Pyridoxine Requirement of the Honey Bee (Apis mellifera) For Brood Rearing". Apidologie. doi:10.1051/apido:19760105.
Jump up ^ Karasov, William H.; Martinez del Rio, Carlos (2008). Physiological Ecology: How Animals Process Energy, Nutrients, and Toxins. Princeton. pp. 63–66.
Jump up ^ Kuhnholz, Susanne (1997). "The Control of Water Collection in Honey Bee Colonies". Behavioral Ecology and Sociobiology. doi:10.1007/s002650050402.
^ Jump up to: a b Kwadha, Charles A.; Ong’amo, George O.; Ndegwa, Paul N.; Raina, Suresh K.; Fombong, Ayuka T. (2017-06-09). "The Biology and Control of the Greater Wax Moth, Galleria mellonella". Insects. 8 (2): 61. doi:10.3390/insects8020061.
Jump up ^ Bryony, Bonning (11 November 2009). "Honey Bee Disease Overview" (PDF). Journal of Invertebrate Pathology. 103: s2-s4. doi:10.1016/j.jip.2009.07.015. Retrieved 21 October 2014.
Jump up ^ McDonald-Gibson, Charlotte. "'Victory for bees' as European Union bans neonicotinoid pesticides blamed for destroying bee population". The Independent. Retrieved 2 July 2014.
Jump up ^ "Colony Collapse Disorder". Beeologics. Archived from the original on 6 February 2013. Retrieved 23 October 2014.
Jump up ^ Crane E (1990). "Honey from honeybees and other insects". Ethology Ecology & Evolution. 3 (sup1): 100–105. doi:10.1080/03949370.1991.10721919.
Jump up ^ Sanford, M.T.; Dietz, A. (1976). "The fine structure of the wax gland of the honey bee (Apis mellifera L.)". Apidologie. 7: 197–207. doi:10.1051/apido:19760301.
Jump up ^ Gillott, Cedric (1995). Entomology. Springer. p. 79.
Jump up ^ Simone-Finstrom, Michael; Spivak, Marla (May–June 2010). "Propolis and bee health: The natural history and significance of resin use by honey bees". Apidologie. 41 (3): 295–311. doi:10.1051/apido/2010016.
Jump up ^ "Propolis:MedlinePlus Supplements". U.S. National Library of Medicine. January 19, 2012.
Jump up ^ Gambichler T; Boms S; Freitag M (April 2004). "Contact dermatitis and other skin conditions in instrumental musicians". BMC Dermatol. 4: 3. doi:10.1186/1471-5945-4-3. PMC 416484  . PMID 15090069.
Jump up ^ Jung-Hoffmann, L (1966). "Die Determination von Königin und Arbeiterin der Honigbiene". Z Bienenforsch. 8: 296–322.
Jump up ^ "Scientific Opinion" (PDF). EFSA Journal. 9 (4): 2083. 2011.
Jump up ^ "Federal Government Seizes Dozens of Misbranded Drug Products: FDA warned company about making medical claims for bee-derived products". Food and Drug Administration. Apr 5, 2010.
Jump up ^ Leung, R; Ho, A; Chan, J; Choy, D; Lai, CK (March 1997). "Royal jelly consumption and hypersensitivity in the community". Clin. Exp. Allergy. 27 (3): 333–6. doi:10.1111/j.1365-2222.1997.tb00712.x. PMID 9088660.
Jump up ^ "Bee castes". Visual Dictionary, QA International. 2017. Retrieved 18 May 2017.
^ Jump up to: a b c d e f g h "Getting Started: Honey Bee Biology". University of Georgia College of Agricultural and Environmental Sciences. 2017. Retrieved 18 May 2017.
^ Jump up to: a b c d e f "Worker, drone and queen bees". PerfectBee LLC. 2017. Retrieved 18 May 2017.
Jump up ^ Harbo JR, Rinderer TE (1980). "Breeding and Genetics of Honey Bees". Beesource Beekeeping. Retrieved 18 May 2017.
Jump up ^ "Morphology of a honeybee: worker". Visual Dictionary, QA International. 2017. Retrieved 18 May 2017.
Jump up ^ Hadisoesilo, Soesilawati. "The Comparative Study of Two Species of Cavity-Nesting Honey Bees of Sulawesi, Indonesia" (PDF).
Jump up ^ C. H. Thawley. "Heat tolerance as a weapon". Davidson College. Archived from the original on October 2, 2013. Retrieved June 1, 2010.
Jump up ^ Michio Sugahara; Fumio Sakamoto (2009). "Heat and carbon dioxide generated by honeybees jointly act to kill hornets". Naturwissenschaften. 96 (9): 1133–6. doi:10.1007/s00114-009-0575-0. PMID 19551367.
Jump up ^ Victoria Gill (July 3, 2009). "Honeybee mobs overpower hornets". BBC News. Retrieved July 5, 2009.
Jump up ^ Giant Honeybees Use Shimmering 'Mexican Waves' To Repel Predatory Wasps - ScienceDaily
Jump up ^ Radloff, Sara E.; Hepburn, H. Randall; Engel, Michael S. (2011). Honeybees of Asia. Berlin: Springer Science & Business Media. ISBN 978-3642164217.
Jump up ^ Hudewenz, Anika; Klein, Alexandra-Maria (2013-12-01). "Competition between honey bees and wild bees and the role of nesting resources in a nature reserve". Journal of Insect Conservation. 17 (6): 1275–1283. doi:10.1007/s10841-013-9609-1. ISSN 1366-638X.
Jump up ^ Johnson, Brian R.; Nieh, James C. (2010-11-01). "Modeling the Adaptive Role of Negative Signaling in Honey Bee Intraspecific Competition". Journal of Insect Behavior. 23 (6): 459–471. doi:10.1007/s10905-010-9229-5. ISSN 0892-7553. PMC 2955239  . PMID 21037953.
Jump up ^ Thompson, Helen; Hunt, Lynn (1999). "Extrapolating from Honeybees to Bumblebees in Pesticide Risk Assessment". Ecotoxicology: 147–166. |access-date= requires |url= (help)
Jump up ^ Tarpy, David (2016). "The Honey Bee Dance Language". NC State Extension.
Jump up ^ Rogers, Lesley J.; Elisa Rigosi; Elisa Frasnelli; Giorgio Vallortigara (27 June 2013). "A right antenna for social behaviour in honeybees". Scientific Reports. Macmillan Publishers. 3. doi:10.1038/srep02045.
Jump up ^ Jessica Shugart. "Honeybees use right antennae to tell friend from foe". Science News. Retrieved 12 March 2016.
Jump up ^ Gorman, James (18 April 2016). "Do Honeybees Feel? Scientists Are Entertaining the Idea" – via NYTimes.com.
Jump up ^ "The symbols of empire". Napoleon.org. Retrieved June 1, 2010.
Jump up ^ "O Asvins, lords of brightness, anoint me with the honey of the bee, that I may speak forceful speech among men! Atharva Veda 91-258, quoted in Maguelonne Toussaint-Samat (Anthea Bell, tr.) The History of Food, 2nd ed. 2009:14.
Jump up ^ Virgil, Georgics, book IV.
Jump up ^ Bee Wilson (2004). The Hive: The Story of the Honeybee. London: John Murray. p. 14. ISBN 0-7195-6598-7.
Jump up ^ "Utah State Motto and Emblem". Retrieved 2017-10-13.
Jump up ^ "Section 1-601". Utah Code 63G General Government. 2017.
Jump up ^ "Section 1-501". Utah Code 63G General Government. 2017.
Jump up ^ "Section 1a-8". Utah Code 67 State Officers and Employees. 2017.
Further reading[edit]
Adam, Brother. In Search of the Best Strains of Bees. Hebden Bridge, W. Yorks: Northern Bee Books, 1983.
Adam, Brother. Bee-keeping at Buckfast Abbey. Geddington, Northants: British Bee Publications, 1975.
Aldersey-Williams, H. Zoomorphic: New Animal Architecture. London: Laurence King Publishing, 2003.
Alexander, P. Rough Magic: A Biography of Sylvia Plath. New York: Da Capo Press, 2003.
Allan, M. Darwin and his flowers. London: Faber & Faber, 1977.
Alston, F. Skeps, their History, Making and Use. Hebden Bridge, W. Yorks: Northern Bee Books, 1987.
Barrett, P. The Immigrant Bees 1788 to 1898, 1995.
Barrett, P. William Cotton.
Beuys, J. Honey is Flowing in All Directions. Heidelberg: Edition Staeck, 1997.
Bevan, E. The Honey-bee: Its Natural History, Physiology and Management. London: Baldwin, Cradock & Joy, 1827.
Bill, L. For the Love of Bees. Newton Abbot, Devon: David & Charles, 1989.
Bodenheimer, F.S. Insects as Human Food The Hague: Dr. W. Junk, 1951.
Brothwell, D., Brothwell, P. Food in Antiquity. London: Thames & Hudson, 1969.
Engel, Michael S. & Grimaldi, David (2005): Evolution of the Insects. Cambridge University Press.
Kak, Subhash C. (1991): The Honey Bee Dance Language Controversy. The Mankind Quarterly Summer 1991: 357–365. HTML fulltext
Lanman, Connor H. The Plight of the Bee: The Ballad of Man and Bee. San Francisco, 2008.
Lindauer, Martin (1971): Communication among social bees. Harvard University Press.
External links[edit]
Wikimedia Commons has media related to Apis (insect).
Look up honey bee in Wiktionary, the free dictionary.
Beediseases Dr. Guido Cordoni's Honey bee Diseases Website
The history of beekeeping
Local Honey for Allergies
Could a Mushroom Save a Honeybee Documentary produced by Oregon Field Guide
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text:
Address Resolution Protocol
From Wikipedia, the free encyclopedia
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v
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e
The Address Resolution Protocol (ARP) is a communication protocol used for discovering the link layer address, such as a MAC address, associated with a given network layer address, typically an IPv4 address. This mapping is a critical function in the Internet protocol suite. ARP was defined in 1982 by RFC 826,[1] which is Internet Standard STD 37.
ARP has been implemented with many combinations of network and data link layer technologies, such as IPv4, Chaosnet, DECnet and Xerox PARC Universal Packet (PUP) using IEEE 802 standards, FDDI, X.25, Frame Relay and Asynchronous Transfer Mode (ATM). IPv4 over IEEE 802.3 and IEEE 802.11 is the most common usage.
In Internet Protocol Version 6 (IPv6) networks, the functionality of ARP is provided by the Neighbor Discovery Protocol (NDP).
Contents
[hide]
1 Operating scope
2 Packet structure
3 Example
4 ARP probe
5 ARP announcements
6 ARP mediation
7 Inverse ARP and Reverse ARP
8 ARP spoofing and Proxy ARP
9 Alternatives to ARP
10 ARP stuffing
11 Standards documents
12 See also
13 References
14 External links
Operating scope[edit]
The Address Resolution Protocol is a request-response protocol whose messages are encapsulated by a link layer protocol. It is communicated within the boundaries of a single network, never routed across internetworking nodes. This property places ARP into the link layer of the Internet protocol suite.[2]
Packet structure[edit]
The Address Resolution Protocol uses a simple message format containing one address resolution request or response. The size of the ARP message depends on the upper layer and lower layer address sizes, which are given by the type of networking protocol (usually IPv4) in use and the type of hardware or virtual link layer that the upper layer protocol is running on. The message header specifies these types, as well as the size of addresses of each. The message header is completed with the operation code for request (1) and reply (2). The payload of the packet consists of four addresses, the hardware and protocol address of the sender and receiver hosts.
The principal packet structure of ARP packets is shown in the following table which illustrates the case of IPv4 networks running on Ethernet. In this scenario, the packet has 48-bit fields for the sender hardware address (SHA) and target hardware address (THA), and 32-bit fields for the corresponding sender and target protocol addresses (SPA and TPA). Thus, the ARP packet size in this case is 28 bytes. The EtherType for ARP is 0x0806. (This appears in the Ethernet frame header when the payload is an ARP packet. Not to be confused with PTYPE below, which appears within this encapsulated ARP packet.)
Internet Protocol (IPv4) over Ethernet ARP packet
octet offset
0
1
0
Hardware type (HTYPE)
2
Protocol type (PTYPE)
4
Hardware address length (HLEN) Protocol address length (PLEN)
6
Operation (OPER)
8
Sender hardware address (SHA) (first 2 bytes)
10
(next 2 bytes)
12
(last 2 bytes)
14
Sender protocol address (SPA) (first 2 bytes)
16
(last 2 bytes)
18
Target hardware address (THA) (first 2 bytes)
20
(next 2 bytes)
22
(last 2 bytes)
24
Target protocol address (TPA) (first 2 bytes)
26
(last 2 bytes)
Hardware type (HTYPE)
This field specifies the network protocol type. Example: Ethernet is 1.
Protocol type (PTYPE)
This field specifies the internetwork protocol for which the ARP request is intended. For IPv4, this has the value 0x0800. The permitted PTYPE values share a numbering space with those for EtherType.[3][4][5]
Hardware length (HLEN)
Length (in octets) of a hardware address. Ethernet addresses size is 6.
Protocol length (PLEN)
Length (in octets) of addresses used in the upper layer protocol. (The upper layer protocol specified in PTYPE.) IPv4 address size is 4.
Operation
Specifies the operation that the sender is performing: 1 for request, 2 for reply.
Sender hardware address (SHA)
Media address of the sender. In an ARP request this field is used to indicate the address of the host sending the request. In an ARP reply this field is used to indicate the address of the host that the request was looking for. (Not necessarily address of the host replying as in the case of virtual media.) Switches do not pay attention to this field, particularly in learning MAC addresses. The ARP PDU is encapsulated in Ethernet frame, and that is what Layer 2 devices examine.
Sender protocol address (SPA)
Internetwork address of the sender.
Target hardware address (THA)
Media address of the intended receiver. In an ARP request this field is ignored. In an ARP reply this field is used to indicate the address of the host that originated the ARP request.
Target protocol address (TPA)
Internetwork address of the intended receiver.
ARP protocol parameter values have been standardized and are maintained by the Internet Assigned Numbers Authority (IANA).[6]
Example[edit]
Two computers in an office (computer 1 and computer 2) are connected to each other in a local area network by Ethernet cables and network switches, with no intervening gateways or routers. Computer 1 has a packet to send to Computer 2. Through DNS, it determines that Computer 2 has the IP address 192.168.0.55. To send the message, it also requires Computer 2's MAC address. First, Computer 1 uses a cached ARP table to look up 192.168.0.55 for any existing records of Computer 2's MAC address (00:eb:24:b2:05:ac). If the MAC address is found, it sends an Ethernet frame with destination address 00:eb:24:b2:05:ac, containing the IP packet onto the link. If the cache did not produce a result for 192.168.0.55, Computer 1 has to send a broadcast ARP message (destination FF:FF:FF:FF:FF:FF MAC address), which is accepted by all computers, requesting an answer for 192.168.0.55. Computer 2 responds with its MAC and IP addresses. Computer 2 may insert an entry for Computer 1 into its ARP table for future use. Computer 1 caches the response information in its ARP table and can now send the packet.[7]
ARP probe[edit]
An ARP probe is an ARP request constructed with an all-zero sender IP address (SPA). The term is used in the IPv4 Address Conflict Detection specification (RFC 5227). Before beginning to use an IPv4 address (whether received from manual configuration, DHCP, or some other means), a host implementing this specification must test to see if the address is already in use, by broadcasting ARP probe packets.[8]
ARP announcements[edit]
ARP may also be used as a simple announcement protocol. This is useful for updating other hosts' mappings of a hardware address when the sender's IP address or MAC address has changed. Such an announcement, also called a gratuitous ARP message, is usually broadcast as an ARP request containing the sender's protocol address (SPA) in the target field (TPA=SPA), with the target hardware address (THA) set to zero. An alternative way is to broadcast an ARP reply with the sender's hardware and protocol addresses (SHA and SPA) duplicated in the target fields (TPA=SPA, THA=SHA).
The gratuitous ARP request message and the gratuitous ARP reply messages are standards-based methods,[9][10] but the "ARP Request" is preferred.[11] Some devices may be configured for the use of either of these two types of GARP.[12]
An ARP announcement is not intended to solicit a reply; instead it updates any cached entries in the ARP tables of other hosts that receive the packet. The operation code may indicate a request or a reply because the ARP standard specifies that the opcode is only processed after the ARP table has been updated from the address fields.[13][14][15]
Many operating systems perform gratuitous ARP during startup. That helps to resolve problems which would otherwise occur if, for example, a network card was recently changed (changing the IP-address-to-MAC-address mapping) and other hosts still have the old mapping in their ARP caches.
Gratuitous ARP is also used by some interface drivers to provide load balancing for incoming traffic. In a team of network cards, it is used to announce a different MAC address within the team that should receive incoming packets.
ARP announcements can be used to defend link-local IP addresses in the Zeroconf protocol (RFC 3927), and for IP address takeover within high-availability clusters.[clarification needed][example needed]
ARP mediation[edit]
ARP mediation refers to the process of resolving Layer 2 addresses through a Virtual Private Wire Service (VPWS) when different resolution protocols are used on the connected circuits, e.g., Ethernet on one end and Frame Relay on the other. In IPv4, each Provider Edge (PE) device discovers the IP address of the locally attached Customer Edge (CE) device and distributes that IP address to the corresponding remote PE device. Then each PE device responds to local ARP requests using the IP address of the remote CE device and the hardware address of the local PE device. In IPv6, each PE device discovers the IP address of both local and remote CE devices and then intercepts local Neighbor Discovery (ND) and Inverse Neighbor Discovery (IND) packets and forwards them to the remote PE device.[16]
Inverse ARP and Reverse ARP[edit]
Inverse Address Resolution Protocol (Inverse ARP or InARP) is used to obtain Network Layer addresses (for example, IP addresses) of other nodes from Data Link Layer (Layer 2) addresses. It is primarily used in Frame Relay (DLCI) and ATM networks, in which Layer 2 addresses of virtual circuits are sometimes obtained from Layer 2 signaling, and the corresponding Layer 3 addresses must be available before those virtual circuits can be used.[17]
Since ARP translates Layer 3 addresses to Layer 2 addresses, InARP may be described as its inverse. In addition, InARP is implemented as a protocol extension to ARP: it uses the same packet format as ARP, but different operation codes.
The Reverse Address Resolution Protocol (Reverse ARP or RARP), like InARP, translates Layer 2 addresses to Layer 3 addresses. However, in ARP the requesting station queries the Layer 3 address of another node, whereas RARP is used to obtain the Layer 3 address of the requesting station itself for address configuration purposes. RARP is obsolete; it was replaced by BOOTP, which was later superseded by the Dynamic Host Configuration Protocol (DHCP).[18]
ARP spoofing and Proxy ARP[edit]
A successful ARP spoofing attack allows an attacker to perform a man-in-the-middle attack.
Main article: ARP spoofing
Main article: Proxy ARP
Because ARP does not provide methods for authenticating ARP replies on a network, ARP replies can come from systems other than the one with the required Layer 2 address. An ARP proxy is a system which answers the ARP request on behalf of another system for which it will forward traffic, normally as a part of the network's design, such as for a dialup internet service. By contrast, in ARP spoofing the answering system, or spoofer, replies to a request for another system's address with the aim of intercepting data bound for that system. A malicious user may use ARP spoofing to perform a man-in-the-middle or denial-of-service attack on other users on the network. Various software exists to both detect and perform ARP spoofing attacks, though ARP itself does not provide any methods of protection from such attacks.[19]
Alternatives to ARP[edit]
Each computer maintains a database of the mapping of Layer 3 addresses (e.g., IP addresses) to Layer 2 addresses (e.g., Ethernet MAC addresses), which is maintained primarily by the reception of ARP packets form the local network link. Thus, it is often called the ARP cache. Traditionally, other methods were also used to maintain this table, such as static configuration files,[20] or centrally maintained lists.
Since at least the 1980s,[21] networked computers have a utility called 'arp' for interrogating or manipulating this table.[22][23][24]
ARP stuffing[edit]
Embedded systems such as networked cameras[25] and networked power distribution devices,[26] which lack a user interface, can use so-called ARP stuffing to make an initial network connection, although this is a misnomer, as ARP is not involved.
This is a solution to an issue in network management of consumer devices, specifically the allocation of IP addresses of ethernet devices where:
the user doesn't have the ability to control DHCP or similar address allocation protocols
the device doesn't have a user interface to configure it with
the user's computer can't communicate with it because it has no suitable IP address.
The solution adopted is as follows:
The user's computer has an IP address stuffed manually into its address table (normally with the arp command with the MAC address taken from a label on the device)
The computer sends special packets to the device, typically a ping packet with a non-default size.
The device then adopts this IP address
The user then communicates with it by telnet or web protocols to complete the configuration.
Such devices typically have a method to disable this process once the device is operating normally, as it is vulnerable to attack.
Standards documents[edit]
RFC 826 - Ethernet Address Resolution Protocol, Internet Standard STD 37.
RFC 903 - Reverse Address Resolution Protocol, Internet Standard STD 38.
RFC 2390 - Inverse Address Resolution Protocol, draft standard
RFC 5227 - IPv4 Address Conflict Detection, proposed standard
See also[edit]
ARP spoofing
Arping
Arptables
Arpwatch
Bonjour Sleep Proxy
Cisco HDLC
Neighbor Discovery Protocol
Proxy ARP
References[edit]
Jump up ^ David C. Plummer (November 1982). "RFC 826, An Ethernet Address Resolution Protocol -- or -- Converting Network Protocol Addresses to 48.bit Ethernet Address for Transmission on Ethernet Hardware". Internet Engineering Task Force, Network Working Group.
Jump up ^ Braden, R. (October 1989). "RFC 1122 - Requirements for Internet Hosts -- Communication Layers". Internet Engineering Task Force.
Jump up ^ IANA ARP - "Protocol Type"
Jump up ^ IANA - Ethertype values
Jump up ^ RFC 5342
Jump up ^ "IANA ARP parameter assignments". IANA. 2009-04-24.
Jump up ^ Chappell, Laura A. and Tittel, Ed. Guide to TCP/IP, Third Edition. Thomson Course Technology, 2007, pp. 115-116.
Jump up ^ Cheshire, S. (July 2008). "RFC 5227 - IPv4 Address Conflict Detection". Internet Engineering Task Force.
Jump up ^ Perkins, C. (November 2010). "RFC 5944 - IP Mobility Support for IPv4, Revised". Internet Engineering Task Force. A gratuitous ARP MAY use either an ARP Request or an ARP Reply packet. [...] any node receiving any ARP packet (Request or Reply) MUST update its local ARP cache with the Sender Protocol and Hardware Addresses in the ARP packet [...]
Jump up ^ Perkins, C. (October 1996). "RFC 2002 - IP Mobility Support". Internet Engineering Task Force.
Jump up ^ Cheshire, S. (July 2008). "RFC 5227 - IPv4 Address Conflict Detection". Internet Engineering Task Force. Why Are ARP Announcements Performed Using ARP Request Packets and Not ARP Reply Packets?
Jump up ^ "FAQ: The Firewall Does not Update the Address Resolution Protocol Table". Citrix. 2015-01-16. [...] garpReply enabled [...] generates ARP packets that [...] are of OPCODE type REPLY, rather than REQUEST.
Jump up ^ Gratuitous ARP in DHCP vs. IPv4 ACD Draft Archived October 12, 2007, at the Wayback Machine.
Jump up ^ RFC 2002 Section 4.6
Jump up ^ RFC 2131 DHCP – Last lines of Section 4.4.1
Jump up ^ Shah, H.; et al. (June 2012). "RFC 6575 Address Resolution Protocol (ARP) Mediation for IP Interworking of Layer 2 VPNs". Internet Engineering Task Force.
Jump up ^ T. Bradley; et al. (September 1998). "RFC 2390 - Inverse Address Resolution Protocol". Internet Engineering Task Force.
Jump up ^ Finlayson, Mann, Mogul, Theimer (June 1984). "RFC 903 - A Reverse Address Resolution Protocol". Internet Engineering Task Force. CS1 maint: Uses authors parameter (link)
Jump up ^ Steve Gibson (2005-12-11). "ARP Cache Poisoning". GRC.
Jump up ^ Sun Microsystems. "SunOS manual page for ethers(5) file". Retrieved 2011-09-28.
Jump up ^ University of California, Berkeley. "BSD manual page for arp(8C) command". Retrieved 2011-09-28.
Jump up ^ Canonical. "Ubuntu manual page for arp(8) command". Archived from the original on 2012-03-16. Retrieved 2011-09-28.
Jump up ^ Apple Computer. "Mac OS X manual page for arp(8) command". Retrieved 2011-09-28.
Jump up ^ Microsoft. "Windows help for arp command". Retrieved 2011-09-28.
Jump up ^ Axis Communication. "Axis P13 Network Camera Series Installation Guide" (PDF). Retrieved 2011-09-28.
Jump up ^ American Power Corporation. "Switched Rack Power Distribution Unit Installation and Quick Start Manual" (PDF). Retrieved 2011-09-28.
This article is based on material taken from the Free On-line Dictionary of Computing prior to 1 November 2008 and incorporated under the "relicensing" terms of the GFDL, version 1.3 or later.
External links[edit]
Wikiversity has learning resources about Address Resolution Protocol
ARP Sequence Diagram (pdf)
Gratuitous ARP
ARP-SK ARP traffic generation tools
Sample Capture file from WireSharkWiki
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