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==Behaviour== The bee-eaters are [[Diurnality|diurnal]], although a few species may migrate during the night if the terrain ''en route'' is unsuitable for stopping or if they are crossing the sea. Bee-eaters are highly social, and pairs sitting or roosting together are often so close that they touch (an [[individual distance]] of zero). Many species are [[bird colony|colonial]] in the breeding season and some species are also highly gregarious when not nesting.<ref name="hbwfamily" /> The social structures of the red-throated bee-eater and the white-fronted bee-eaters have been described as more complex than for any other bird species.<ref name="hbwfamily" /> The birds exist in colonies located on nesting cliffs, and have a stable structure all year round. These colonies typically contain five to 50 burrows, occasionally up to 200, and are composed of clans of two or three pairs, their [[Helpers at the nest|helpers]], and their offspring.<ref name="emlen" /> The helpers are male offspring from a previous year.<ref name="hbwfamily" /> Within the colony, the males alternate between guarding their mate and attempting to make forced copulations with other females.<ref name="emlen" /> The females in turn attempt to lay eggs in their neighbour's nests, an example of [[Brood parasite|brood parasitism]]. Some individuals also specialise in [[kleptoparasitism]], stealing prey collected by other colony members. The colony's daily routine is to emerge from the nesting holes or roosting branches soon after dawn, preen and sun themselves for an hour, then disperse to feed. Feeding territories are divided by clan, with each clan defending its territory from all others of the same species, including clans of the same colony.<ref name="hed" /> The clans return to the colony before dusk, and engage in more social behaviour before retiring for the night. Colonies are situated several hundred metres apart and have little to do with each other, although young individuals may disperse between colonies. As such, these species can be thought to have four tiers of social kinship: the individual pair, the family unit, the clan, and the colony as a whole.<ref name="hbwfamily" /> Bee-eaters spend around 10% of their day on comfort activities. These include [[sun bathing|sunning themselves]], [[dust bath]]ing and water bathing. Sunning behaviour helps warm birds in the morning, reducing the need to use energy to raise their temperature. It also has a social aspect, as multiple birds adopt the same posture. Finally, it may help stimulate [[parasite]]s in the feathers, making them easier to find and remove. Due to their hole-nesting lifestyle, bee-eaters accumulate a number of external parasites such as [[mite]]s and flies. Together with sunning, bouts of dust bathing (or water bathing where available), as well as rigorous [[preening]], keep the feathers and skin in good health. Bathing with water involves making shallow dives into a water body and then returning to a perch to preen.<ref name="hbwfamily" /> ===Diet and feeding=== [[File:Blue-throated Bee-eater Courtship offering.jpg|thumb|left|A male [[blue-throated bee-eater]] presents his mate with a captured insect]] The bee-eaters are almost exclusively aerial hunters of insect prey. Prey is caught either on the wing or more commonly from an exposed perch from which the bee-eater watches for prey. Smaller, rounder-winged bee-eaters typically hunt from branches and twigs closer to the ground, whereas the larger species hunt from tree tops or telephone wires. One unusual technique often used by carmine bee-eaters is to ride on the backs of [[bustard]]s.<ref name="hbwfamily" /> Prey can be spotted from a distance; European bee-eaters are able to spot a bee {{convert|60|m|ft|abbr=on}} away, and blue-cheeked bee-eaters have been observed flying out {{convert|100|m|ft|abbr=on}} to catch large wasps. Prey is approached directly or from behind. Prey that lands on the ground or on plants is usually not pursued. Small prey may be eaten on the wing, but larger items are returned to the perch where they are beaten until dead and then broken up. Insects with poisonous stings are first smacked on the branch, then, with the bird's eyes closed, rubbed to discharge the [[venom]] sac and [[stinger]]. This behaviour is [[Intrinsic and extrinsic properties|innate]], as demonstrated by a juvenile bird in captivity, which performed the task when first presented with wild bees. This bird was stung on the first five tries, but by ten bees, it was as adept at handling bees as adult birds.<ref name="hbwfamily" /> Bee-eaters consume a wide range of insects; beyond a few distasteful butterflies they consume almost any insect from tiny ''[[Drosophila]]'' flies to large beetles and dragonflies. At some point bee-eaters have been recorded eating beetles, [[mayfly|mayflies]], [[stonefly|stoneflies]], [[cicada]]s, [[termite]]s, [[Orthoptera|crickets and grasshopper]]s, [[mantis]]es, true flies and moths. For many species, the dominant prey item are stinging members of the order Hymenoptera, namely [[wasp]]s and [[bee]]s. In a survey of 20 studies, the proportion of the diet made up by bees and wasps varied from 20% to 96%, with the average being 70%. Of these [[honeybee]]s can comprise a large part of the diet, as much as 89% of the overall intake. The preference for bees and wasps may have arisen because of the numerical abundance of these suitably sized insects.<ref name="hbwfamily" /> The [[Apis dorsata|giant honeybee]] is a particularly commonly eaten species. These bees attempt to congregate in a mass defence against the bee-eaters.<ref name="kastberger" /> In Israel, a European bee-eater was documented attempting to eat a small bat that it had caught, which probably could not fit down its throat.<ref>{{cite magazine|last=Sarchet |first=Penny |url=https://www.newscientist.com/article/dn27818-ambitious-bee-eater-attempts-to-swallow-a-bat-whole/ |title=Ambitious bee-eater attempts to swallow a bat whole |magazine=New Scientist |date=2015-07-01 |access-date=2017-06-29}}</ref> Like kingfishers, bee-eaters regurgitate [[Pellet (ornithology)|pellets]] of undigested material, typically {{convert|2|cm|in|sigfig=1|abbr=on}} long black oblongs.<ref name="hbwfamily" /> ==== Predation of honey bees ==== [[File:Colonie de prigorii (Merops apiaster) vandalizată de apicultori.jpg|thumb|right|Bee-eater colony destroyed by bee-keepers. The entrances into the bee eater's nests were deliberately blocked with stones]] If an [[apiary]] is set up close to a bee-eater colony, a larger number of honey bees are eaten because they are more abundant. However, studies show the bee-eaters do not intentionally fly into the apiary, rather they feed on the insects caught on pastures and meadows within a radius of {{convert|12|km|mi|abbr=on}} from the colony, this maximum distance being reached only when there is a shortage of food. Observations show that the birds actually enter the apiary only in cold and rainy periods, when the bees do not leave the hive and other insect prey are harder for the bee-eaters to detect.<ref>{{cite web|url=https://sor.ro/ro/noutati/Prigonirea-prigoriei-Mituri-si-adevaruri-despre-albine-si-albinarel.html|title=Prigonirea prigoriei. [Myths and truths about honey bees and bee eaters ]|publisher=Romanian Ornithological Society|language=ro|access-date=2018-06-27|archive-date=2018-06-27|archive-url=https://web.archive.org/web/20180627144415/https://sor.ro/ro/noutati/Prigonirea-prigoriei-Mituri-si-adevaruri-despre-albine-si-albinarel.html|url-status=dead}}</ref> Many [[bee-keeper]]s believe that the bee-eaters are the main obstacle causing worker bees not to forage, and instead stay inside the hives for much of the day between May and the end of August. However, a study carried out in a [[eucalyptus]] forest in the Alaluas region in the [[Murqub District]] in Libya, {{convert|80|km|mi|abbr=on}} east of [[Tripoli, Libya|Tripoli]], showed that the bee-eaters were not the main obstacle to bee foraging; in some cases, the foraging rate was higher in the presence of the birds than in their absence. The average bird meal consisted of 90.8% honey bees and 9.2% beetles.<ref>{{cite web|last=Alfallah|first=H.M|url=http://www.moa.gov.cy/moa/da/ead/ead.nsf/0/6722A01C440AA5EDC2257B1D003949BC/$file/Bee-eaters-Bee-Keepers.pdf|title=The impact of the Bee-eater ''Merops apiaster'' on the behavior of honey bee ''Apis mellifera'' L. during foraging|publisher=Mansoura Journal of Plant Protection and Pathology, 1(12): 1023–1030|access-date=2018-06-27|archive-url=https://web.archive.org/web/20180627144323/http://www.moa.gov.cy/moa/da/ead/ead.nsf/0/6722A01C440AA5EDC2257B1D003949BC/$file/Bee-eaters-Bee-Keepers.pdf|archive-date=2018-06-27|url-status=dead}}</ref> Predation is more likely when the bees are queening or during the peak of migration, from late March till mid-April, and in mid-September. Hives close to or under trees or overhead cables are at increased risk as the birds pounce on flying insects from these perches.<ref name=toM >{{cite news | last = Carabott| first = Sarah| title =Bee-eater is not to blame for decline in honey bees | newspaper = Times of Malta| location = Valletta| date = 2015-10-26| url = https://www.timesofmalta.com/articles/view/20151026/local/bee-eater-is-not-to-blame-for-decline-in-honey-bees.589691| access-date = 2018-06-27}}</ref> ===Breeding=== [[File:מצוק השרקרקים.JPG|thumb|left|Bee-eater nesting cliff in [[Modi'in]], [[Israel]]]] Bee-eaters are [[monogamy in animals|monogamous]] during a nesting season, and in sedentary species, pairs may stay together for multiple years. Migratory bee-eaters may find new mates each breeding season. The courtship displays of the bee-eaters are rather unspectacular, with some calling and raising of throat and wing feathers. The exception is the performance of the white-throated bee-eater. Their "butterfly display" involves both members of a pair performing a gliding display flight with shallow wing-beats; they then perch facing each other, raising and folding their wings while calling.<ref name="hbwfamily" /> Most members of the family engage in courtship feeding, where the male presents prey items to the female, and such feeding can account for much, if not all, of the energy females require for egg creation.<ref name="akh" /> Like almost all Coraciiformes the bee-eaters are cavity [[bird nest|nesters]].<ref name="eberh" /> In the case of the bee-eaters the nests are burrows dug into the ground, either into the sides of earth cliffs or directly into level soil. Both types of nesting site are vulnerable, those on level ground are vulnerable to trampling and small predators, whereas those in cliffs, which are often the banks of rivers, are vulnerable to flash floods, which can wipe out dozens or hundreds of nests. Many species will nest either on cliffs or on level ground but prefer cliffs, although [[Böhm's bee-eater]] always nests on level ground. The burrows are dug by both birds in the pair, sometimes assisted by helpers. The soil or sand is loosened with jabs of the sharp bill, then the feet are used to kick out the loose soil. It has been suggested that riverine [[loess]] deposits that do not crumble when excavated may be favoured by the larger bee-eaters.<ref name="loessbee" /><ref name="loessc" /> There may be several false starts where nests are dug partway before being abandoned; in solitary species this can give the impression of colonial living even when that is not the case. The process of nest building can take as long as twenty days to complete, during which time the bill can be blunted and shortened. Nests are generally used only for a single season and are rarely used twice by the bee-eaters, but abandoned nests may be used by other birds, snakes and bats as shelter and breeding sites.<ref name="ecosystem" /> No nesting material is used in the breeding cavity.<ref name="frybeeater19"/> One white egg is laid each day until the typical clutch of about five eggs is complete.<ref name="hbwfamily" /> [[egg incubation|Incubation]] starts soon after the first egg is laid, with both parents sharing this duty in the day, but only the female at night. The eggs hatch in about 20 days, and the newly hatched young are blind, pink and naked. For most species, the eggs do not all hatch at the same time, so if food is in short supply only the older chicks survive.<ref name="hbwfamily" /> Adults and young [[defecation|defecate]] in the nest, and their pellets are trodden underfoot, making the nest cavity very malodorous.<ref name="frybeeater19"/> The chicks are in the nest for about 30 days.<ref name="hbwfamily" /> Bee-eaters may nest as single pairs, loose colonies or dense colonies. Smaller species tend to nest solitarily, while medium-sized bee-eaters have small colonies, and larger and migratory species nest in large colonies that can number in the thousands. In some instances, colonies may contain more than one species of bee-eater.<ref name="kf98" /> In species that nest gregariously, breeding pairs may be assisted by up to five helpers.<ref name="ffh" /> These birds may alternate between breeding themselves and helping in successive years.<ref name="hbwfamily" />
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