Editing Hummingbird (section)

=== Song, vocal learning, and hearing ===
[[File:Acoustic-Divergence-with-Gene-Flow-in-a-Lekking-Hummingbird-with-Complex-Songs-pone.0109241.s010.oga|thumb|Complex songs of male [[wedge-tailed sabrewing]] hummingbirds (''Campylopterus curvipennis'') in [[lek mating|mating leks]] of eastern Mexico<ref name="Gonz">{{cite journal | last1=González | first1=Clementina | last2=Ornelas | first2=Juan Francisco | title=Acoustic divergence with gene flow in a lekking hummingbird with complex songs | journal=PLOS ONE| volume=9 | issue=10 | date=2014-10-01 | issn=1932-6203 | doi=10.1371/journal.pone.0109241 | page=e109241|pmid=25271429|pmc=4182805 | bibcode=2014PLoSO...9j9241G | doi-access=free }}</ref>]]

Many hummingbird species exhibit a diverse vocal repertoire of chirps, squeaks, whistles and buzzes.<ref name="duque">{{cite journal |author1=Duque, F.G. |author2=Carruth, L.L. |title=Vocal communication in hummingbirds |journal=Brain, Behavior and Evolution |volume=97 |issue=3–4 |pages=241–252 |date=2022 |pmid=35073546 |doi=10.1159/000522148 |s2cid=246278322 |url=https://www.karger.com/Article/FullText/522148|type=Review|doi-access=free }}</ref><ref name="clo">{{Cite web |date=2015 |title=Song sounds of various hummingbird species |url=https://www.allaboutbirds.org/guide/browse.aspx?shape=37,11 |access-date=25 June 2016 |website=All About Birds |publisher=The Cornell Lab of Ornithology, Cornell University, Ithaca, New York}}</ref> Vocalizations vary in complexity and spectral content during social interactions, foraging, territorial defense, courtship, and mother-nestling communication.<ref name=duque/> Territorial vocal signals may be produced in rapid succession to discourage aggressive encounters, with the chirping rate and loudness increasing when intruders persist.<ref name=duque/> During the breeding season, male and female hummingbirds vocalize as part of courtship.<ref name=duque/>

Hummingbirds exhibit vocal production learning to enable song variation {{ndash}} "dialects" that exist across the same species.<ref name=duque/> For example, the blue-throated hummingbird's song differs from typical oscine songs in its wide frequency range, extending from 1.8&nbsp;kHz to about 30&nbsp;kHz.<ref name="pytte">{{Cite journal |last1=Pytte |first1=C.L. |last2=Ficken |first2=M.S. |last3=Moiseff |first3=A. |year=2004 |title=Ultrasonic singing by the blue-throated hummingbird: A comparison between production and perception |url=https://www.researchgate.net/publication/8542654 |journal=Journal of Comparative Physiology A |volume=190 |issue=8 |pages=665–673 |doi=10.1007/s00359-004-0525-4 |pmid=15164219 |s2cid=7231117}}</ref> It also produces [[ultrasound|ultrasonic]] vocalizations which do not function in communication.<ref name="pytte"/> As blue-throated hummingbirds often alternate singing with catching small flying insects, it is possible the ultrasonic clicks produced during singing disrupt insect flight patterns, making insects more vulnerable to predation.<ref name="pytte"/> Anna's, Costa's, long-billed hermits, and Andean hummingbirds have song dialects that vary across habitat locations and phylogenetic clades.<ref name=duque/><ref name="duque2">{{cite journal |author1=Duque, F.G. |author2=Rodríguez-Saltos, C.A. |author3=Wilczynsk, W. |title=High-frequency vocalizations in Andean hummingbirds |journal=Current Biology |volume=28 |issue=17 |pages=R927–R928 |date=September 2018 |pmid=30205060 |doi=10.1016/j.cub.2018.07.058 |s2cid=52188456 |doi-access=free |bibcode=2018CBio...28.R927D }}</ref>
[[File:Calypte anna - Anna's Hummingbird XC109651.mp3|thumb|Song of male Anna's hummingbird (''Calypte anna'')]]

The avian vocal organ, the [[Syrinx (bird anatomy)|syrinx]], plays an important role in understanding hummingbird song production.<ref name="Monte2020">{{Cite journal |last1=Monte |first1=Amanda |last2=Cerwenka |first2=Alexander F. |last3=Ruthensteiner |first3=Bernhard |last4=Gahr |first4=Manfred |last5=Düring |first5=Daniel N. |date=2020-07-06 |title=The hummingbird syrinx morphome: a detailed three-dimensional description of the black jacobin's vocal organ |journal=BMC Zoology |volume=5 |issue=1 |pages=7 |doi=10.1186/s40850-020-00057-3 |issn=2056-3132 |doi-access=free |s2cid=220509046|hdl=20.500.11850/429165 |hdl-access=free }}</ref> What makes the hummingbird's syrinx different from that of other birds in the Apodiformes order is the presence of internal muscle structure, accessory cartilages, and a large [[Eardrum|tympanum]] that serves as an attachment point for external muscles, all of which are adaptations thought to be responsible for the hummingbird's increased ability in pitch control and large frequency range.<ref name="Monte2020"/><ref name="Riede2020">{{Cite journal |last1=Riede |first1=Tobias |last2=Olson |first2=Christopher R. |date=2020-02-06 |title=The vocal organ of hummingbirds shows convergence with songbirds |journal=Scientific Reports |language=en |volume=10 |issue=1 |pages=2007 |bibcode=2020NatSR..10.2007R |doi=10.1038/s41598-020-58843-5 |issn=2045-2322 |pmc=7005288 |pmid=32029812}}</ref>

Hummingbird songs originate from at least seven specialized [[nucleus (neuroanatomy)|nuclei]] in the [[forebrain]].<ref name="jarvis">{{Cite journal |last1=Jarvis |first1=Erich D. |last2=Ribeiro |first2=Sidarta |last3=da Silva |first3=Maria Luisa |last4=Ventura |first4=Dora |last5=Vielliard |first5=Jacques |last6=Mello |first6=Claudio V. |year=2000 |title=Behaviourally driven gene expression reveals song nuclei in hummingbird brain |journal=Nature |volume=406 |issue=6796 |pages=628–632 |bibcode=2000Natur.406..628J |doi=10.1038/35020570 |pmc=2531203 |pmid=10949303}}</ref><ref>{{Cite journal |last=Gahr M. |year=2000 |title=Neural song control system of hummingbirds: comparison to swifts, vocal learning (Songbirds) and nonlearning (Suboscines) passerines, and vocal learning (Budgerigars) and nonlearning (Dove, owl, gull, quail, chicken) nonpasserines |journal=J Comp Neurol |volume=486 |issue=2 |pages=182–196 |doi=10.1002/1096-9861(20001016)426:2<182::AID-CNE2>3.0.CO;2-M |pmid=10982462 |s2cid=10763166}}</ref> A [[genetic expression]] study showed that these nuclei enable [[vocal learning]] (ability to acquire vocalizations through imitation), a rare trait known to occur in only two other groups of birds ([[parrot]]s and [[songbird]]s) and a few groups of mammals (including humans, [[cetacea|whales and dolphins]], and [[bat]]s).<ref name="jarvis"/> Within the past 66 million years, only hummingbirds, parrots, and songbirds out of 23 bird [[order (biology)|orders]] may have independently evolved seven similar forebrain structures for singing and vocal learning, indicating that evolution of these structures is under strong [[epigenetics|epigenetic]] constraints possibly derived from a common ancestor.<ref name="jarvis"/><ref>{{Cite journal |last1=Renne |first1=Paul R. |last2=Deino |first2=Alan L. |last3=Hilgen |first3=Frederik J. |last4=Kuiper |first4=Klaudia F. |last5=Mark |first5=Darren F. |last6=Mitchell |first6=William S. |last7=Morgan |first7=Leah E. |last8=Mundil |first8=Roland |last9=Smit |first9=Jan |display-authors=3 |date=7 February 2013 |title=Time Scales of Critical Events Around the Cretaceous-Paleogene Boundary |url=http://www.cugb.edu.cn/uploadCms/file/20600/20131028144132060.pdf |url-status=dead |journal=Science |volume=339 |issue=6120 |pages=684–687 |bibcode=2013Sci...339..684R |doi=10.1126/science.1230492 |pmid=23393261 |archive-url=https://web.archive.org/web/20170207164818/http://www.cugb.edu.cn/uploadCms/file/20600/20131028144132060.pdf |archive-date=7 February 2017 |access-date=1 April 2018 |s2cid=6112274}}</ref>

Generally, birds have been assessed to vocalize and hear in the range of 2–5&nbsp;kHz, with hearing sensitivity falling with higher frequencies.<ref name=duque2/> In the [[Ecuadorian hillstar]] (''Oreotrochilus chimborazo''), vocalizations were recorded in the wild to be at a frequency above 10&nbsp;kHz, well outside of the known hearing ability of most birds.<ref name=duque2/> Song system nuclei in the hummingbird brain are similar to those songbird brains, but the hummingbird brain has specialized regions involved for song processing.<ref name=duque/>