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Greater honeyguide
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==Behaviour and ecology== The greater honeyguide is known to guide people to the nests of wild bees.<ref name=IsackReyer1989>{{cite journal |author1=Isack, H. A. |author2=H.-U. Reyer |title=Honeyguides and honey gatherers: interspecific communication in a symbiotic relationship |journal=Science |volume=243 |year=1989 |pages=1343–1346 |doi=10.1126/science.243.4896.1343 |pmid=17808267 |issue=4896 |bibcode=1989Sci...243.1343I |s2cid=4220280}}</ref><ref name=Wood2014>{{Cite journal |last1=Wood |first1=B. M.|last2=Pontzer|first2=H. |last3=Raichlen|first3=D. A. |last4=Marlowe|first4=F. W. |date=2014 |title=Mutualism and manipulation in Hadza–honeyguide interactions |journal=Evolution and Human Behavior |volume=35 |issue=6|pages=540–546 |doi=10.1016/j.evolhumbehav.2014.07.007 |bibcode=2014EHumB..35..540W |url=https://www.sciencedirect.com/science/article/pii/S1090513814000877|url-access=subscription }}</ref> A guiding bird attracts a person's attention with wavering, chattering {{"'}}tya' notes compounded with peeps or pipes",<ref>{{cite book |author1=Short, L. |author2=Horne, J. |year=2002 |editor1=Del Hoyo, J. |editor2=Elliott, A. |editor3=Sargatal, J. |title=Handbook of the Birds of the World: Jacamars to Woodpeckers |volume=7 |chapter=Family Indicatoridae (Honeyguides) |chapter-url=http://www.hbw.com/hbw/volume7/family7.html |publisher=Lynx Edicions |isbn=978-84-87334-37-5 |title-link=Handbook of the Birds of the World}}</ref> sounds it also gives in aggression. The guiding bird flies toward an occupied nest (greater honeyguides know the sites of many bees' nests in their territories) and then stops nearby the nest. Honey-hunters then do a final search for the bee colony, and if deemed suitable, harvest honey from the bee colony through the use of fire and smoke to subdue the bees, and axes and machetes to expose the colony. After harvesting the honey, the honeyguide eats wax that is left.<ref name="ShortHorneDiamond2003">{{cite book |author1=Short, L. |author2 =Horne, J. |author3=Diamond, A. W. |name-list-style=amp |year=2003 |title=Firefly Encyclopedia of Birds |chapter=Honeyguides |editor=Perrins, C. |editor-link=Christopher Perrins |pages=396–397 |publisher=Firefly Books |isbn=978-1-55297-777-4 |chapter-url-access=registration |chapter-url =https://archive.org/details/fireflyencyclope0000unse/page/396}}</ref> One study found that use of honeyguides by the [[Borana Oromo|Boran people]] of East Africa reduces their search time for honey by approximately two-thirds. Because of this benefit, the Boran use a specific loud whistle, known as the ''fuulido'', when a search for honey is about to begin. The ''fuulido'' doubles the encounter rate with honeyguides.<ref name=IsackReyer1989/> In northern Tanzania, Honeyguides increased the Hadza's rate of finding bee nests by 560%, and led men to significantly higher yielding nests than those found without honeyguides.<ref name=Wood2014/> Another study of the [[Yao people (East Africa)|Yao]] honey-hunters in northern Mozambique showed that the honeyguides responded to the traditional ''brrrr-hmm'' call of the honey-hunters. The chances of finding a bee-hive were greatly increased when the traditional call was used. That study reported anecdotes from Yao honey-hunters that adult but not juvenile honeyguides respond to the specific honey-hunting calls.<ref>{{cite journal |journal=Science |year=2016 |title=Reciprocal signaling in honeyguide-human mutualism |author1=Spottiswoode, C. N. |author1-link=Claire Spottiswoode |author2=Begg, K. S.|author3=Begg, C. M. |volume=353 |pages=387–389 |issue=6297 |url=https://www.repository.cam.ac.uk/handle/1810/256963|access-date=2020-08-14 |doi=10.1126/science.aaf4885 |pmid=27463674|bibcode=2016Sci...353..387S |s2cid=206648494}}</ref> In African folklore, it is frequently noted that the honeyguide should be thanked with a gift of honey; if not, it may lead its follower to a [[lion]], bull [[elephant]], or venomous [[snake]] as punishment. However, “others maintain that honeycomb spoils the bird, and leave it to find its own bits of comb”.<ref name=ShortHorneDiamond2003/> While many depictions of the human-honeyguide mutualism emphasize honey-hunters graciously repaying the birds with piles of wax left in a conspicuous location, such behavior is not universal. The Hadza people of northern Tanzania frequently burn, bury, or hide the wax that lays with the intent of keeping the bird hungry, and more likely to guide again.<ref name=Wood2014/> Some greater honeyguides have stopped this guiding behavior, or mutualism, in parts of Kenya, and this is thought to be due to a loss of response from people in the area.<ref>{{Cite journal |last1=Dean |first1=W. R. J. |last2=Siegfried |first2=W. R. |last3=MacDonald|first3=I. A. W. |date=1990 |title=The Fallacy, Fact, and Fate of Guiding Behavior in the Greater Honeyguide |jstor=2385968|journal=Conservation Biology |volume=4 |issue=1|pages=99–101 |doi=10.1111/j.1523-1739.1990.tb00272.x|bibcode=1990ConBi...4...99D}}</ref><ref>{{Cite journal |last=Friedmann|first=H. |date=1955 |title=The Honey-Guides |journal=United States National Museum Bulletin |volume=208 |page=50 |url=https://library.si.edu/digital-library/book/bulletinunitedst2081955unit}}</ref> A study done in 2025 found that sometimes honeyguides do lead humans to animals that are not bees, such as snakes. However the researches think that this is probably due to error by the birds and not punitive behavior.<ref>{{Cite journal |last=Lloyd-Jones |first=David J. |last2=Muamedi |first2=Musaji |last3=Spottiswoode |first3=Claire N. |date=2025 |title=To Bees or Not to Bees: Greater Honeyguides Sometimes Guide Humans to Animals Other Than Bees, but Likely Not as Punishment |url=https://onlinelibrary.wiley.com/doi/10.1002/ece3.71136 |journal=Ecology and Evolution |language=en |volume=15 |issue=4 |pages=e71136 |doi=10.1002/ece3.71136 |issn=2045-7758 |pmc=12037988 |pmid=40303557}}</ref> === Possible guiding of animals === {{multiple image |direction=vertical |align=right |image1=Indicator indicator 1838, crop.jpg |caption1=Adult male illustrated by [[Nicolas Huet the Younger|Nicolas Huet]] |image2=Indicator indicator, 2023-05-20 07h07, 26s gidsroep te Kamp Rynoue naby Pretoria, a.mp3 |caption2=Guiding call near [[Pretoria]], South Africa}} Many sources say that this species also guides [[honey badger]]s.<ref>{{cite book |author=Attenborough, D. |year=1998 |title=The Life of Birds |url= https://archive.org/details/lifeofbirds0000atte |url-access=registration |publisher=Princeton University Press |isbn=978-0-691-01633-7 |author-link=David Attenborough}}</ref><ref>{{cite book |author=Estes, R. D. |year=1999 |title=The Safari Companion: A Guide to Watching African Mammals |publisher=Chelsea Green |pages=361–362 |isbn=978-1-890132-44-6 |url=https://books.google.com/books?id=Xqp7poFviNcC}}</ref><ref>{{cite book |author1=Zimmerman, D. A. |author2=Turner, D. A. |author3=Pearson, D. J. |name-list-style=amp |year=1999 |title=Birds of Kenya and Northern Tanzania |publisher=Princeton University Press |page=406 |isbn=978-0-691-01022-9}}</ref> [[Anders Sparrman|Sparrman]] noted in the 18th century that indigenous Africans reported this interaction, but Friedmann adds that no biologist has seen it. Friedmann quotes reports that greater honeyguides guide [[baboon]]s and speculates that the behavior [[evolution|evolved]] in relation to these species before the appearance of humanity.<ref>{{cite journal |author1=Dean, W. R. J. |author2=MacDonald, I. A. W. | title=A review of African birds feeding in association with mammals |journal=Ostrich |volume=52 |issue=3 |year=1981 |pages=135–155 |doi=10.1080/00306525.1981.9633599|bibcode=1981Ostri..52..135D}}</ref> However, they state, <blockquote>In addition to that listed by Friedmann (1955:41-47), the only recent record is of a greater honeyguide giving its guiding call to baboons at Wankie Game Reserve, Zimbabwe (C. J. Vernon, pers. comm.). However, Vernon did not see a positive response by the baboons to the honeyguide. No additional records of honeyguides and ratels have been reported since Friedmann (1955) and the first-hand accounts given in his review in support of this association are all of incomplete guiding sequences. No biologist has ever reported this association.</blockquote> Honeyguides are thought to guide other animals, a behavior that may have evolved with "early human". Later studies estimate that interaction between honeyguides and honey badgers likely occurs, "but is highly localized or extremely difficult to observe, or both".<ref>{{cite journal|author1=Van der Wal, J. E. M. |author2=Afan, A. I.|author3=Anyawire, M. |author4=Begg, C. M.|author5=Begg, K. S.|author6=Dabo, G. A.|author7=Gedi, I. I.|author8=Harris, J. A.|author9=Isack, H. A. |author10=Ibrahim, J. I.|author11=Jamie, G. A.|author12=Kamboe, W.-B. W.|author13=Kilawi, A. O.|author14=Kingston, A.|author15=Laltaika, E. A. |author16=Lloyd-Jones, D. J.|author17=M'manga, G. M.|author18=Muhammad, N. Z.|author19=Ngcamphalala, C. A.|author20=Nhlabatsi, S. O. |author21=Oleleteyo, T. T. |author22=Sanda, M.|author23=Tsamkxao, L.|author24=Wood, B. M.|author25=Spottiswoode, C. N.|author26=Cram, D. L. |title=Do honey badgers and greater honeyguide birds cooperate to access bees' nests? Ecological evidence and honey-hunter accounts |journal=Journal of Zoology |volume=321|issue=1|year=2023 |pages=22–32 |doi=10.1111/jzo.13093 |doi-access=free|url=https://www.repository.cam.ac.uk/bitstreams/30809086-febc-475c-aad2-6db6184213ab/download}}</ref> It has also been acknowledged that bee colonies are seasonally very common in Africa and ratels probably have no trouble finding them.<ref>{{cite book |author1=Short, L. |author2=Horne, J. |author2-link=Jennifer F. M. Horne |name-list-style=amp |title=Toucans, Barbets and Honeyguides |pages=473–480 |publisher=Oxford University Press |year=2002 |isbn=978-0-19-854666-5}}</ref> Another argument against guiding of non-human animals is that near cities, where Africans increasingly buy [[sugar]] rather than hunting for wild honey, guiding behavior is disappearing. Ultimately, it may disappear everywhere.<ref name=ShortHorneDiamond2003 /> ===Diet=== [[File:Greater Honeyguide, Gambia.jpg|thumb|Immature male in [[The Gambia]]]] {{Unreferenced section|date=January 2023}} The greater honeyguide feeds primarily on the contents of [[bee]] colonies: [[beeswax]], [[egg (biology)|eggs]], [[larva]]e and [[pupa]]e; and [[waxworm]]s. It enters bees' nests while the bees are torpid in the early morning, feeds at abandoned hives and scavenges at hives robbed by people or other large animals, notably the honey badger. It frequently associates with other honeyguides at bees' nests; immatures dominate adults, and immatures of this species dominate all others. ===Reproduction=== The greater honeyguide is a [[brood parasite]]. It lays white eggs in series of 3 to 7, for a total of 10 to 20 in a year. Each egg is laid in a different nest of a bird of another species, including some woodpeckers, [[Lybiidae|barbets]], [[kingfisher]]s, [[bee-eater]]s, [[wood hoopoe]]s, [[starling]]s, and large [[hirundinidae|swallows]]. It is common for the female greater honeyguide to break the host's eggs when laying her own.<ref>{{Cite journal |last1=Spottiswoode |first1=C. N. |last2=Colebrook-Robjent |first2=J. F. R. |date=2007 |title=Egg puncturing by the brood parasitic Greater Honeyguide and potential host counteradaptations |journal=Behavioral Ecology |volume=18|issue=4 |pages=792–799 |doi=10.1093/beheco/arm025 |doi-access=free |hdl=10.1093/beheco/arm025 |hdl-access=free}}</ref> All the species parasitized nest in holes, covered nests, or deep cup nests. The chick has a membranous hook on the bill that it uses, while still blind and featherless, to kill the host's young outright or by repeated wounds.
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