Editing Bee-eater

{{short description|Widespread group of insectivorous bird species in the family Meropidae }}
{{For|the family of scorpionflies|Meropeidae}}
{{featured article}}
{{Automatic taxobox
| image             = African bee-eaters composite.jpg
| image_caption     = Six common African bee-eaters
| taxon             = Meropidae
| authority         = [[Constantine Samuel Rafinesque-Schmaltz|Rafinesque]], 1815
| subdivision_ranks = Genera
| subdivision       =
* ''[[Nyctyornis]]''
* ''[[Purple-bearded bee-eater|Meropogon]]''
* ''[[Merops (genus)|Merops]]''
| range_map         = Meropidaemap2.png
| range_map_caption =
<div style="text-align:left;"><big>{{Legend2|#FF0000| Approximate area in which <br />bee-eater species regularly breed | border=1px solid #aaa}}</big></div>
}}
The '''bee-eaters''' are a group of [[bird]]s in the [[family (biology)|family]] '''Meropidae''', containing three genera and thirty-one species. Most species are found in Africa and Asia, with a few in southern Europe, Australia, and New Guinea. They are characterised by richly coloured [[plumage]], slender bodies, and usually elongated central tail feathers. All have long down-turned [[Beak|bills]] and medium to long wings, which may be pointed or round. Male and female plumages are usually similar.

As their name suggests, bee-eaters predominantly eat flying [[insect]]s, especially [[bee]]s and [[wasp]]s, which are caught on the wing from an open perch. The insect's [[stinger]] is removed by repeatedly hitting and rubbing the insect on a hard surface. During this process, pressure is applied to the insect's body, thereby discharging most of the [[venom]].

Most bee-eaters are [[Social animal|gregarious]]. They form [[Bird colony|colonies]], nesting in burrows tunnelled into vertical sandy banks, often at the side of a river or in flat ground. As they mostly live in colonies, large numbers of nest holes may be seen together. The eggs are white, with typically five to the clutch. Most species are [[monogamy in animals|monogamous]], and both parents care for their young, sometimes with assistance from related birds in the colony.

Bee-eaters may be killed by [[bird of prey|raptors]]; their nests are raided by rodents, weasels, martens and snakes, and they can carry various [[parasite]]s. Some species are adversely affected by human activity or [[Habitat destruction|habitat loss]], but none meet the [[International Union for Conservation of Nature]]'s vulnerability criteria, and all are therefore evaluated as "[[Least-concern species|least concern]]". Their conspicuous appearance means that they have been mentioned by ancient writers and incorporated into [[mythology]].

==Taxonomy==
[[File:Rainbow Bee-eaters Juffs.JPG|thumb|[[Rainbow bee-eater]]s, a ''[[Merops (genus)|Merops]]'' species]]
The bee-eaters were first named as a scientific group by the French [[polymath]] [[Constantine Samuel Rafinesque|Constantine Samuel Rafinesque-Schmaltz]], who created the bird subfamily Meropia for these birds in 1815.<ref name="raff"/><ref name="bock"/> The name, now modernised as Meropidae, is derived from ''Merops'', the [[Ancient Greek]] for "bee-eater",<ref name="job"/> and the English term "bee-eater" was first recorded in 1668, referring to the European species.<ref name="oed"/>

The bee-eaters have been considered to be related to other families, such as the [[Coraciidae|roller]]s, [[hoopoe]]s and [[kingfisher]]s, but ancestors of those families diverged from the bee-eaters at least forty million years ago, so any relationship is not close.<ref name="hbwfamily" /> The scarcity of [[fossil]]s is unhelpful. Bee-eater fossils from the [[Pleistocene]] (2,588,000 to 11,700 years ago) have been found in Austria, and there are [[Holocene]] (from 11,700 years ago to present) specimens from Israel and Russia, but all have proved to be of the extant [[European bee-eater]].<ref name="frybeeater195"/> Opinions have varied as to the bee-eater's nearest relatives. In 2001, Fry considered the kingfishers to be the most likely,<ref name="hbwfamily" /> whereas a large study published in 2008 found that bee-eaters are [[sister group|sister]] to all other [[Coraciiformes]] (rollers, [[ground roller]]s, [[tody|todies]], [[motmot]]s and kingfishers).<ref name= hackett/> A 2009 book supported Fry's contention,<ref name= mayr/> but then a later study in 2015 suggested that the bee-eaters are sister to the rollers.<ref name= prum/> The 2008 and 2015 papers both linked the kingfishers to the New World motmots.<ref name= hackett/><ref name= prum/>

More recent [[molecular phylogenetic]] studies have confirmed that the bee-eaters are more closely related to the rollers and ground rollers than they are to the todies, motmots and kingfishers. The relationship between the families is shown the cladogram below.<ref>{{Cite journal | last1=Kuhl | first1=H. | last2=Frankl-Vilches | first2=C. | last3=Bakker | first3=A. | last4=Mayr | first4=G. | last5=Nikolaus | first5=G. | last6=Boerno | first6=S.T. | last7=Klages | first7=S. | last8=Timmermann | first8=B. | last9=Gahr | first9=M. | date=2021 | title=An unbiased molecular approach using 3′-UTRs resolves the avian family-level tree of life | journal=Molecular Biology and Evolution | volume=38 | issue=1 | pages=108–127 | doi=10.1093/molbev/msaa191 | doi-access=free | pmid=32781465 | pmc=7783168 | hdl=21.11116/0000-0007-B72A-C | hdl-access=free }}</ref><ref>{{ cite journal | last1=Stiller | first1=J. | display-authors=etal | year=2024 | title=Complexity of avian evolution revealed by family-level genomes | journal=Nature | volume=629 | issue= 8013| pages=851–860 | doi=10.1038/s41586-024-07323-1 | doi-access=free | pmid=38560995 | pmc=11111414 | bibcode=2024Natur.629..851S }}</ref> The number of species in each family is taken from the list maintained by [[Frank Gill (ornithologist)|Frank Gill]], [[Pamela C. Rasmussen]] and David Donsker on behalf of the [[International Ornithological Committee]] (IOC).<ref name=ioc>{{cite web| editor1-last=Gill | editor1-first=Frank | editor1-link=Frank Gill (ornithologist) | editor2-last=Donsker | editor2-first=David | editor3-last=Rasmussen | editor3-first=Pamela | editor3-link=Pamela C. Rasmussen | date=December 2023 | title=IOC World Bird List Version 14.1 | url=https://www.worldbirdnames.org/new/ | publisher=International Ornithologists' Union | access-date=17 June 2024 }}</ref>
{{Clade | style=font-size:100%;line-height:100%
|label1=Coraciiformes
|1={{clade
   |1={{clade
      |1='''Meropidae''' – bee-eaters (31 species)
      |2={{clade
         |1=[[Brachypteraciidae]] – ground rollers (5 species)
         |2=[[Coraciidae]] – rollers (13 species)
         }}
      }}
   |2={{clade
      |1=[[Todidae]] – todies (5 species)
      |2={{clade
         |1=[[Momotidae]] – motmots (14 species)
         |2=[[Alcedinidae]] – kingfishers (118 species)
         }}
      }}
   }}
}}

The bee-eaters are generally similar in appearance, although they are normally divided into three [[genus|genera]]. ''[[Nyctyornis]]'' comprises two large species with long throat feathers, the [[blue-bearded bee-eater]] and the [[red-bearded bee-eater]], both of which have rounded wings, a ridged [[culmen (bird)|culmen]], feathered nostrils and a relatively sluggish lifestyle. The [[purple-bearded bee-eater]] is the sole member of ''Meropogon'', which is intermediate between ''Nyctyornis'' and the typical bee-eaters, having rounded wings and a "beard", but a smooth culmen and no nostril feathers. All the remaining species are normally retained in the single genus ''[[Merops (genus)|Merops]]''. There are close relationships within this genus, for example the [[red-throated bee-eater]] and the [[white-fronted bee-eater]] form a [[superspecies]], but formerly suggested genera, such as ''Aerops'', ''Melittophagus'', ''Bombylonax'' and ''Dicrocercus'',<ref name=peters/> have not been generally accepted for several decades since a 1969 paper united them in the current arrangement.<ref name="hbwfamily" /><ref name="fry69"/>

===Species in taxonomic order===
{{cladogram|title=Evolutionary relationships
|caption=[[Phylogenetic tree]] (maximum parsimony) based on a 2007 study. ''Nyctyornis athertoni'' and ''Merops revoilii'' were not included in the study. The placement of ''Meropogon'' is unclear.<ref name="phylo" />
|cladogram=
{{clade|style=font-size:75%;line-height:75%
 |1=''Nyctyornis''
 |2={{clade
  |1={{clade
   |1=''Merops bullockoides''
   |2=''M. bulocki''
  }}
  |2={{clade 
   |1={{clade
    |state1=dashed
    |1=''Meropogon forsteni''
    |2=''Merops breweri''
   }}
   |2={{clade
    |1={{clade
     |1=''M. gularis''
     |2=''M. muelleri''
    }}
    |2={{clade
     |1=''M. hirundineus''
     |2={{clade
      |1=''M. oreobates''
      |2={{clade
       |1=''M. pusillus''
       |2=''M. variegatus''
      }}
     }}
    }}
   }}
   |3={{clade
    |1={{clade
     |1={{clade
      |1=''M. boehmi''
     }}
     |2={{clade
      |1=''M. albicollis''
      |2={{clade
       |1=''M. nubicus''
       |2={{clade
        |1=''M. malimbicus''
        |2={{clade
         |1={{clade
          |1=''M. orientalis''
          |2={{clade
           |1=''M. leschenaulti''
           |2=''M. viridis''
          }}
         }}
         |2={{clade
          |1=''M. philippinus''
          |2={{clade
           |1={{clade
            |1=''M. apiaster''
            |2=''M. ornatus''
           }}
           |2={{clade
            |1=''M. persicus''
            |2=''M. superciliosus''
           }}
          }}
         }}
        }}
       }}
      }}
     }}
    }}
   }}
  }}
 }}
}}
}}

The bee-eater family contains the following species.
{| class="wikitable"
|-
! Image !! Genus   !! Living Species
|-
|[[File:Nyctyornis amictus - Kaeng Krachan.jpg|175px]]||'''''[[Nyctyornis]]''''' {{small|Jardine & Selby, 1830}}||
* [[Red-bearded bee-eater]], ''Nyctyornis amictus''
* [[Blue-bearded bee-eater]], ''Nyctyornis athertoni''
|-
|[[File:Purple-bearded Bee-eater - Sulawesi MG 5287.jpg|175px]]||'''''Meropogon''''' {{small|Bonaparte, 1850}}||
* [[Purple-bearded bee-eater]], ''Meropogon forsteni''
|-
|[[File:Merops bulocki frenatus.jpg|175px]]||'''''[[Merops (genus)|Merops]]''''' {{small|Linnaeus, 1758}}||
* [[Little bee-eater]], ''Merops pusillus''
* [[Blue-cheeked bee-eater]], ''Merops persicus''
* [[Green bee-eater|Asian green bee-eater]], ''Merops orientalis''
*[[African green bee-eater]], ''Merops viridissimus''
*[[Arabian green bee-eater]], ''Merops cyanophyrs''
* [[White-throated bee-eater]], ''Merops albicollis''
* [[Swallow-tailed bee-eater]], ''Merops hirundineus''
* [[Blue-tailed bee-eater]], ''Merops philippinus''
* [[Black bee-eater]], ''Merops gularis''
* [[Blue-headed bee-eater]], ''Merops muelleri''
* [[Blue-moustached bee-eater]], ''Merops mentalis''
* [[Red-throated bee-eater]], ''Merops bulocki''
* [[White-fronted bee-eater]], ''Merops bullockoides''
* [[Blue-breasted bee-eater]], ''Merops variegatus''
* [[Ethiopian bee-eater]], ''Merops lafresnayii''
* [[Cinnamon-chested bee-eater]], ''Merops oreobates''
* [[Black-headed bee-eater]], ''Merops breweri''
* [[Somali bee-eater]], ''Merops revoilii''
* [[Böhm's bee-eater]], ''Merops boehmi''
* [[Blue-throated bee-eater]], ''Merops viridis''
* [[Rufous-crowned bee-eater]], ''Merops americanus''
* [[Olive bee-eater]], ''Merops superciliosus''
* [[Rainbow bee-eater]], ''Merops ornatus''
* [[European bee-eater]], ''Merops apiaster''
* [[Chestnut-headed bee-eater]], ''Merops leschenaulti''
* [[Rosy bee-eater]], ''Merops malimbicus''
* [[Northern carmine bee-eater]], ''Merops nubicus''
* [[Southern carmine bee-eater]], ''Merops nubicoides''
|-
|}

The Asian green bee-eater, African green bee-eater, and Arabian green bee-eater were previously considered to be a single species, and are still treated as such by some authorities.<ref name="hbwasianb" /><ref name="hbwarabb" /><ref name="hbwafricanb" />

A 2007 [[nuclear DNA|nuclear]] and [[mitochondrial DNA]] study produced a possible [[phylogenetic tree]], although the position of the purple-bearded bee-eater seems anomalous, in that it appears amongst ''Merops'' species.<ref name="phylo" />

==Description==
[[File:Merops bullockoides 1 Luc Viatour.jpg|thumb|''Merops'' species such as the [[white-fronted bee-eater]] usually have a black bar through the eye.]]
The bee-eaters are [[Morphology (biology)|morphologically]] a fairly uniform group. They share many features with related Coraciiformes such as the kingfishers and rollers, being large-headed (although less so than their relatives), short-necked, brightly plumaged and short-legged. Their wings may be rounded or pointed, with the wing shape closely correlated with the species' preferred foraging habitat and [[bird migration|migratory]] tendencies. Shorter, rounder wings are found on species that are sedentary and make typically short foraging flights in denser forests and reed-beds. Those with more elongated wings are more migratory. All the bee-eaters are highly aerial; they take off strongly from perches, fly directly without undulations, and are able to change direction quickly, although they rarely hover.<ref name="hbwfamily" />

The [[flight feather]]s of the wing comprise 10 [[flight feather#primaries|primaries]], the outermost being very small, and 13 [[flight feather#secondaries|secondaries]], and there are 12 [[flight feather#rectrices|tail feathers]].<ref name="frybeeater29"/>

The [[beak|bill]]s of bee-eaters are curved, long and end in a sharp point. The bill can bite strongly, particularly at the tip, and it is used as a pair of [[forceps]] with which to snatch insects from the air and crush smaller prey. The short legs have weak feet, and when it is moving on the ground a bee-eater's gait is barely more than a shuffle. The feet have sharp claws used for perching on vertical surfaces and also for nest excavation.<ref name="hbwfamily" />

The [[plumage]] of the family is generally very bright and in most species is mainly or at least partially green, although the two [[carmine bee-eater (disambiguation)|carmine bee-eaters]] are primarily rose-coloured. Most of the ''Merops'' bee-eaters have a black bar through the eye and many have differently coloured throats and faces. The extent of the green in these species varies from almost complete in the [[green bee-eater]] to barely any green in the [[white-throated bee-eater]]. Three species, from equatorial Africa, have no green at all in their plumage, the [[black bee-eater]], the [[blue-headed bee-eater]] and the [[rosy bee-eater]]. Many species have elongated central tail feathers.<ref name="hbwfamily" />

There is little visible [[sexual dimorphism|difference between the sexes]] in most of the family, although in several species the [[Iris (anatomy)|iris]] is red in the males and brown-red in the females, and in species with tail-streamers these may be slightly longer in males. Both the European and red-bearded bee-eaters have sex-based differences in their plumage colour, and the female rainbow bee-eater has shorter tail streamers than the male, which terminate in a club-shape that he lacks.<ref name="hbwfamily" /> There may be instances where bee-eaters are sexually [[dichroism|dichromatic]] at the [[ultraviolet]] part of the colour spectrum, which humans cannot see. A study of [[blue-tailed bee-eater]] found that males were more colourful than females in UV light. Their overall colour was also affected by body condition, suggesting that there was a signalling component to plumage colour.<ref name="swwy" /> Juveniles are generally similar to adults, except for the two ''Nyctyornis'' species, in which the young have mainly green plumage.<ref name="hbwfamily" />

Bee-eaters have calls that are characteristic for each species. Most sound simple to the human ear, but show significant variability when studied in detail, carrying significant information for the birds.<ref name="hbwfamily" />

==Distribution and habitat==
[[File:Blue-bearded_bee-eater_in_Thailand.jpg|thumb|upright|The [[blue-bearded bee-eater]] is associated with forests, where it forages in edge habitats]]
The bee-eaters have an [[Old World]] distribution, occurring from Europe to Australia. The centre of diversity of the family is Africa, although a number of species also occur in Asia. Single species occur in each of Europe, (the [[European bee-eater]]), Australia (the [[rainbow bee-eater]]) and Madagascar (the [[olive bee-eater]], also found on mainland Africa). Of the three genera, ''Merops'', which has the majority of the species, occurs across the entirety of the family's distribution. ''Nyctyornis'' is restricted to Asia, ranging from India and southern China to the Indonesian islands of [[Sumatra]] and [[Borneo]]. The genus ''Meropogon'' has a single species restricted to [[Sulawesi]] in Indonesia.<ref name="hbwfamily" />

Bee-eaters are fairly indiscriminate in their choice of habitat. Their requirements are simply an elevated perch from which to watch for prey and a suitable ground substrate in which to dig their breeding burrow. Because their prey is entirely caught on the wing they are not dependent on any vegetation type. A single species, the blue-headed bee-eater, is found inside closed rainforest where it forages close to the ground in poor light in the gaps between large trees. Six other species are also closely associated with rainforest, but occur in edge habitat such as along rivers, in tree-fall gaps, off trees overhanging ravines or on emergent tree crowns above the main canopy.<ref name="hbwfamily" />

Species that breed in subtropical or temperate areas of Europe, Asia and Australia are all migratory. The European bee-eaters that breed in southern Europe and Asia migrate to [[West Africa|West]] and [[southern Africa]]. Another population of the same species breeds in South Africa and Namibia; these birds move northwards after breeding. In Australia the rainbow bee-eater is migratory in the southern areas of its range, migrating to Indonesia and New Guinea, but occurs year-round in northern Australia. Several species of bee-eater, are intra-African migrants;<ref name="hbwfamily" /> the white-throated bee-eater, for example, breeds on the southern edge of the [[Sahara]] and winters further south in equatorial [[rainforest]].<ref name="hbwwtbe" /> The most unusual migration is that of the [[southern carmine bee-eater]], which has a three-stage migration; after breeding in a band between Angola and Mozambique it moves south to Botswana, Namibia and South Africa before moving north to its main wintering grounds in northern Angola, Congo and Tanzania.<ref name="carmine migration" />

==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" />

==Predators and parasites==
[[File:Green Bee-eaters (Merops orientalis) at Sindhrot near Vadodara, Gujrat Pix 254.jpg|thumb|[[Green bee-eater|Asian green bee-eater]]s [[dust bath]]ing to keep down parasites]]

Bee-eater nests may be raided by rats and snakes,<ref name="frybeeater" /> and the adults are hunted by birds of prey such as the [[Levant sparrowhawk]].<ref name="christie" /> The little bee-eater and red-throated bee-eaters are hosts of the [[greater honeyguide]] and the [[lesser honeyguide]], both [[brood parasite]]s. The young honeyguides kill the bee-eater's chicks and destroy any eggs. The begging call of the honeyguide sounds like ''two'' bee-eater chicks, ensuring a good supply of food from the adult bee-eaters.<ref name="frybeeater" /><ref name="stab" />

Bee-eaters may be infested by several blood-feeding [[fly|flies]] of the genus ''[[Carnus (fly)|Carnus]]'',<ref name="carnus" /> and the biting fly ''[[Ornithophila|Ornithophila metallica]]''.<ref name="frybeeater" /> Other parasites include [[Mallophaga|chewing lice]] of the genera ''[[Meromenopon]]'', ''[[Brueeliaa]]'' and ''[[Meropoecus]]'', some of which are specialist parasites of bee-eaters,<ref name="saudi" /><ref name="hungary" /> and the stickfast flea ''[[Echidnophaga gallinacea]]''. The hole-nesting lifestyle of bee-eaters means that they tend to carry a higher burden of external parasites than non-hole-nesting bird species.<ref name="frybeeater" /> Bee-eaters may also be infected by [[protozoa]]n blood parasites of the genus ''[[Haemoproteus]]'' including ''[[Haemoproteus meropis|H. meropis]]''.<ref name="iraq" />

Fly [[larva]]e of the genus ''[[Fannia (fly)|Fannia]]'' live in the nests of at least European bee-eaters, and feed on faeces and food remains. Their presence and cleaning activities appear to benefit the developing bee-eaters.<ref name= fannia>{{cite journal | last1 = Krištofík | first1 = Ján | last2 = Darolová | first2 = Alžbeta | last3 = Hoi  | first3 = Christine |last4= Hoi |first4=  Herbert |year =2016| title  = Housekeeping by lodgers: the importance of bird nest fauna on offspring condition | journal = Journal of Ornithology | volume = 158| pages = 245–252 | doi =10.1007/s10336-016-1384-9| s2cid = 43638499 }}</ref>

==Status==
[[File:Southern Carmine Bee-eater (Merops nubicoides) (16732824032).jpg|thumb|The [[southern carmine bee-eater]] is adversely affected by persecution and habitat loss.]]

The [[International Union for Conservation of Nature]] (IUCN) assesses species vulnerability in terms of total population and the rate of any population decline. None of the bee-eaters meet the IUCN vulnerability criteria, and all are therefore evaluated as "[[Least-concern species]]".<ref name="iucnwhole" />

Open country species, which comprise the majority of bee-eaters, have mostly expanded in range as more land is converted to agriculture, but some tropical forest species have suffered declines through loss of habitat, although no species or subspecies gives serious cause for concern. There is some human persecution of bee-eaters, with nest holes being blocked, adults shot or [[birdlime|limed]], or young taken for food. More generally problematic is the unintended destruction of nests. This can occur through cattle trampling, as with the blue-headed bee-eater in Kenya, or loss of forests, with massive conversion of native forest to [[oil palm]] plantations in Malaysia being particularly concerning.<ref name="hbwfamily" />

A study of the southern carmine bee-eater in Zimbabwe showed that it was affected by deliberate interference and persecution and loss of woodlands, and that nesting sites are lost through poor water management leading to river bank damage, dam construction and panning for gold. Colonies are becoming concentrated into the [[national park]]s and the [[Zambezi]] Valley. The well-studied European bee-eater is trapped and shot on migration in countries bordering the [[Mediterranean]], an estimated 4,000–6,000 annually being killed in Cyprus alone, but with a global population of between 170,000 and 550,000 pairs even losses on that scale make little overall impact.<ref name="hbwfamily" />

==In culture==
[[File:Kopf des Aristoteles 1.jpg|thumb|[[Aristotle]] advised the killing of bee-eaters to protect [[beehive|hives]]]]

Bee-eaters were mentioned by ancient writers such as [[Aristotle]] and [[Virgil]], who both advised [[beekeeper]]s to kill the birds. Aristotle knew that bee-eaters nested at the end of tunnels up to {{convert|2|m|ft|abbr=on}} long and the size of their clutch. He said that nesting adults were fed by their own young, based on the observed actual help at the nest by related birds.<ref name="cocker" />

In [[Greek mythology]], the [[Thebes, Greece|Theban]] [[Botres]] was fatally struck by his father when he desecrated a ritual sacrifice of a ram to the god [[Apollo]] by tasting the victim's brains. The god took pity on him, turning him into a bee-eater.<ref name="forbes" />

The [[Ancient Egypt]]ians believed that bee-eaters had medical properties, prescribing the application of bee-eater fat to deter biting flies, and treating the eyes with the smoke from charred bee-eater legs to cure an unspecified female complaint.<ref name="cocker" />

In [[Hinduism]], the shape of the bird in flight was thought to resemble a bow, with the long bill as an arrow. This led to a [[Sanskrit]] name meaning "[[Vishnu]]'s bow" and an association with archer gods. [[Defamation|Scandalmonger]]s were thought to be [[reincarnation|reincarnated]] as bee-eaters, because of the metaphorical poison they bore in their mouths.<ref name="cocker" />

Depictions in classical art are rare for such striking birds. The only known Ancient Egyptian example is a relief, probably of a little green bee-eater, on a wall of Queen [[Hatshepsut|Hatshepsut's]] [[mortuary temple]], and an early [[Ancient Rome|Roman]] mural depicting blue-cheeked bee-eaters was found in the villa of [[Agrippina the Younger|Agrippina]]. Bee-eaters have been depicted on the postage stamps of at least 38 countries, the European and Carmine bee-eaters being the most common subjects, with 18 and 11 countries respectively.<ref name="hbwfamily" />
{{clear}}

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<ref name="raff">{{cite book |last = Rafinesque |first = Constantine Samuel | title = Analyse de la nature: ou, Tableau de l'univers et des corps organisés |volume = 1815 |publisher = Self-published |year = 1815 |location = Palermo |page = 66| language = fr | url =https://www.biodiversitylibrary.org/page/48310144 }}</ref>
<ref name="saudi">{{cite journal |last1=El-Ahmed |first1=A. |last2=Gamal |first2=el-D. N. |last3=Shobrak |first3=M. |last4=Dik |first4=B. |year=2012 |title=First records of the chewing lice (Phthiraptera) associated with European bee eater (''Merops apiaster'') in Saudi Arabia |journal=Journal of the Egyptian Society of Parasitology |volume=42 |issue=3 |pages=525–533 |pmid=23469628 |doi=10.12816/0006338|doi-access=free }}</ref>
<ref name="stab">{{cite journal |last1=Spottiswoode |first1=Claire N. |last2=Koorevaar |first2=Jeroen |title=A stab in the dark: chick killing by brood parasitic honeyguides |journal=  Biology Letters|volume=8 |issue=2 |pages=1–4 |year=2011 |doi=10.1098/rsbl.2011.0739|pmid=21900311 |pmc=3297377 }}</ref>
<ref name="swwy">{{cite journal |last1=Siefferman |first1=Lynn |last2=Wang |first2=Yuan-Jyun |last3=Wang |first3=Yi-Ping |last4=Yuan |first4=Hsiao-Wei |year=2007 |title=Sexual dichromatism, dimorphism, and condition-dependent coloration in blue-tailed bee-eaters |journal=Condor |volume=109 |issue=3 |pages=577–584 |doi=10.1650/8201.1 |s2cid=53383771 |doi-access=free }}</ref>
}}

==External links==
{{Commons category|Meropidae}}
{{Wikispecies|Meropidae}}
* [http://www.hbw.com/ibc/family/bee-eaters-meropidae Bee-eater videos] on the Internet Bird Collection
* [http://montereybay.com/creagrus/bee-eaters.html Meropidae, Bird families of the World]
* [http://tolweb.org/Meropidae/26433 Meropidae on Tree of Life Web]

{{Bee-eaters}}
{{Taxonbar|from=Q183147}}
{{Authority control}}

[[Category:Meropidae|*]]
[[Category:Taxa named by Constantine Samuel Rafinesque]]