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{{Short description|Vocalization used to attract mates}}
{{about|sexual reproduction|other uses|Mating call (disambiguation)}}
 
A '''mating call''' is the auditory signal used by animals to attract mates. It can occur in males or females, but literature is abundantly favored toward researching mating calls in malesfemales. In addition, mating calls are often the subject of [[mate choice]], in which the preferences of one gender for a certain type of mating call can drive [[sexual selection]] in a species. This can result in [[sympatric speciation]] of some animals, where two species diverge from each other while living in the same environment.
 
There are many different mechanisms to produce mating calls, which can be broadly categorized into vocalizations and mechanical calls. Vocalizations are considered as sounds produced by the [[larynx]] and are often seen in species of birds, mammals, amphibians, and insects. Mechanical calls refer to any other type of sound that the animal produces using unique body parts and/or tools for communication with potential mates. Examples include crickets that vibrate their wings, birds that flap their feathers, and frogs that use an air sac instead of lungs.
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=== Birds ===
[[File:Song Sparrow (8381415679).jpg|thumb|Song sparrow]]
[[File:SND 4458a.ogg|thumb|Mating call of Japanese bush warbler, ''Horornis diphone'']]The use of vocalizations is widespread in avian species and are often used to attract mates. Different aspects and features of bird song such as structure, amplitude and frequency have evolved as a result of sexual selection.<ref name=nemeth2012>{{cite journal |last=Nemeth |first=E. |author2=Kempenaers, B. |author3=Matessi, G. |author4= Brumm, H. |title=Rock sparrow song reflects male age and reproductive success |journal=PLOS ONE |year=2012 |volume=7 |issue=8 |pages=e43259 |pmid=22927955 |doi=10.1371/journal.pone.0043259 |pmc=3426517|bibcode=2012PLoSO...743259N |doi-access=free }}</ref><ref>{{cite journal |author1= Mikula, P. |author2= Valcu, M. | author3= Brumm, H. | author4= Bulla, M. | author5= Forstmeier, W. | author6= Petrusková, T. | author7= Kempenaers, B. | author8= Albrecht, T | year=2021| title= A global analysis of song frequency in passerines provides no support for the acoustic adaptation hypothesis but suggests a role for sexual selection. | journal= Ecology Letters | volume= 24 | issue = 3 | pages= 477–486|doi=10.1111/ele.13662|pmid= 33314573|s2cid= 229176172 | doi-access= free |bibcode= 2021EcolL..24..477M }}</ref>
 
Large song repertoires are preferred by females of many avian species.<ref name=pfaff2007>{{cite journal|last=Pfaff|first=J. A. |author2=Zanette, L. |author3=MacDougall-Shackleton, S. A. |author4= MacDougall-Shackleton, E. A. |title=Song repertoire size varies with HVC volume and is indicative of male quality in song sparrows (Melospiza melodia) |journal=[[Proceedings of the Royal Society B]]|date=Aug 22, 2007|volume=274|issue=1621|pages=2035–40 |pmid=17567560|doi=10.1098/rspb.2007.0170 |pmc=2275172}}</ref> One hypothesis for this is that song repertoire is positively correlated with the size of the brain's [[HVC (avian brain region)|song control nucleus]] (HVC). A large HVC would indicate developmental success. In [[song sparrow]]s, males with large repertoires had larger HVCs, better body condition and lower heterophil-to-lymphocyte ratios indicating better immune health. This supports the idea that song sparrows with large song repertoires have better lifetime fitness and that song repertoires are honest indicators of the male's "quality". Possible explanations for this adaptation include direct benefits to the female, such as superior parental care or territory defense, and indirect benefits, such as good genes for their offspring.<ref name=pfaff2007 />
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[[Japanese bush warbler]] songs from island populations have an acoustically simple structure when compared to mainland populations.<ref>{{cite journal |last=Hamao|first=Shoji |title=Acoustic structure of songs in island populations of the Japanese bush warbler, Cettia diphone, in relation to sexual selection |journal=Journal of Ethology|year=2012|volume=31|issue=1|pages=9–15|doi=10.1007/s10164-012-0341-1|doi-access=free}}</ref> Song complexity is correlated with higher levels of sexual selection in mainland populations, showing that a more complex song structure is advantageous in an environment with high levels of sexual selection. Another example is in [[purple-crowned fairywren]]s; larger males of this species sing advertising songs at a lower frequency than smaller rival males. Since body size is a characteristic of good health, lower frequency calls are a form of honest signaling. Negative correlation between body size and call frequency is supported across multiple species within the taxa.<ref>{{cite journal|last=Hall|first=ML|author2=Kingma, SA |author3=Peters, A |title=Male songbird indicates body size with low-pitched advertising songs.|journal=PLOS ONE|year=2013|volume=8|issue=2|pages=e56717|pmid=23437221|doi=10.1371/journal.pone.0056717|pmc=3577745|bibcode=2013PLoSO...856717H|doi-access=free}}</ref> In the [[rock sparrow]], song frequency is positively associated with reproductive success. Slower song rate is associated with age and is preferred by females. Reproductive status of the individual is communicated through higher maximum frequency. There was also positive correlation between age and [[extra-pair copulation]] frequency.<ref name=nemeth2012/>[[File:Red deer stag 2009 denmark.jpg|thumb|Red deer stag during [[rut (mammalian reproduction)|rut]]]]
 
Bird calls are also known to continue after pair formation in several socially [[Monogamous pairing in animals|monogamous]] bird species. In one experimental population of [[zebra finch]]es, there was increased singing activity by the male after breeding.<ref name="bolund2012">{{cite journal|last=Bolund|first=Elisabeth|author2=Schielzeth, Holger |author3=Forstmeier, Wolfgang |title=Singing activity stimulates partner reproductive investment rather than increasing paternity success in zebra finches|journal=Behavioral Ecology and Sociobiology|year=2012|volume=66|issue=6|pages=975–984|doi=10.1007/s00265-012-1346-z|bibcode=2012BEcoS..66..975B |s2cid=17518228}}</ref> This increase is positively correlated with the partner's reproductive investment. The female finches were bred in cages with two subsequent males that differed with varying amounts of song output. Females produced larger [[Bird egg|eggs]] with more orange yolks when paired with a male with a high song output. This suggests that the relative amount of song production in paired zebra finch males might function to stimulate the partner rather than to attract extra-pair females.<ref name="bolund2012" />
 
=== Mammals ===
During the [[breeding season]], mammals will call out to the opposite sex. Male [[koala]]s that are bigger will let out a different sound than smaller koalas. The bigger males which are routinely sought out for are called sires. Females choose sires because of indirect benefits that their offspring could inherit, like larger bodies.<ref>{{cite journal | last1 = Charlton | first1 = B. D. | last2 = Ellis | last3 = Brumm | first3 = J. | last4 = Nilsson | first4 = K. | last5 = Fitch | first5 = W. T. | year = 2012 | title = Female koalas prefer bellows in which lower formants indicate larger males | journal = Animal Behaviour | volume = 84 | issue = 6| pages = 1565–1571 | doi=10.1016/j.anbehav.2012.09.034| s2cid = 53175246 }}</ref> Non-sires and females do not vary in their body mass and can reject a male by screaming or hitting him. Male-male competition is rarely exhibited in koalas.<ref>{{cite journal | last1 = Ellis | first1 = W. A. H. | last2 = Bercovitch | first2 = F. B. | year = 2011 | title = Body size and sexual selection in the koala | journal = Behavioral Ecology and Sociobiology | volume = 65 | issue = 6| pages = 1229–1235 | doi=10.1007/s00265-010-1136-4| bibcode = 2011BEcoS..65.1229E | s2cid = 26046352 }}</ref>
Acoustic signaling is a type of call that can be used from a significant distance encoding an organism's location, condition and identity.<ref>{{cite journal | last1 = Wilkins | first1 = M. R. | last2 = Seddon | first2 = N. | last3 = Safran | first3 = R. J. | year = 2013 | title = Evolutionary divergence in acoustic signals: causes and consequences | journal = Trends in Ecology & Evolution | volume = 28 | issue = 3| pages = 156–66 | doi=10.1016/j.tree.2012.10.002| pmid = 23141110 | bibcode = 2013TEcoE..28..156W }}</ref> [[Sac-winged bat]]s display acoustic signaling, which is often interpreted as songs. When females hear these songs, named a 'whistle', they call onto the males to breed with a screech of their own. This action is termed 'calling of the sexes'.<ref>{{cite journal | year = 2008 | title = Songs, Scents, and Senses: Sexual Selection in the Greater Sac-Winged Bat, Saccopteryx bilineata | journal = Journal of Mammalogy| volume = 89 | issue = 6| pages = 1401–1410 | doi=10.1644/08-mamm-s-060.1| last1 = Voigt | first1 = Christian C. | last2 = Behr | first2 = Oliver | last3 = Caspers | first3 = Barbara | last4 = von Helversen | first4 = Otto | last5 = Knörnschild | first5 = Mirjam | last6 = Mayer | first6 = Frieder | last7 = Nagy | first7 = Martina | doi-access = free }}</ref> [[Red deer]] and [[spotted hyena]]s along with other mammals also perform acoustic signaling.<ref>Logan, C. J. & Clutton-Brock, T. H. [https://www.repository.cam.ac.uk/bitstream/handle/1810/261261/Logan_and_Clutton-Brock-2012-Behavioural_Processes-AM.pdf?sequence=1 Validating methods for estimating endocranial volume in individual red deer (Cervus elaphus)]. ''Behavioural processes'' 2013; 92, 143–6</ref><ref>{{cite journal | last1 = Goller | first1 = K. V | last2 = Fickel | first2 = J. | last3 = Hofer | first3 = H. | last4 = Beier | first4 = S. | last5 = East | first5 = M. L. | year = 2013 | title = Coronavirus genotype diversity and prevalence of infection in wild carnivores in the Serengeti National Park, Tanzania | journal = Archives of Virology | volume = 158 | issue = 4| pages = 729–34 | doi=10.1007/s00705-012-1562-x | pmid=23212740| pmc = 7086904 | doi-access = free }}</ref>
[[File:Tungara Frog (Engystomops pustulosus) Calling.jpg|thumb|Tungara frog]]
 
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Most frogs use an air sac located under their mouth to produce mating calls. Air from the lungs channels to the air sac to inflate it, and the air sac resonates to produce a mating call. The larynx is larger and more developed in males, which causes their call to be louder and stronger<ref>{{cite journal | last1 = Wilczynski | first1 = W. | last2 = McClelland | first2 = B. E. | last3 = Rand | first3 = A. S. | year = 1993 | title = Acoustic, auditory, and morphological divergence in three species of neotropical frog | journal = Journal of Comparative Physiology A | volume = 172 | issue = 4| pages = 425–438 | doi=10.1007/bf00213524| pmid = 8315606 | s2cid = 21923928 }}</ref>
 
In the [[Tungara frog|túngara frog]], males use a whining call followed by up to seven clucks. Males who have a whine-cluck call are more successful in attracting females than males whose call is a whine alone. The ability to produce clucks is due to a specialized fibrous mass attached to the frog's vocal folds, creating an unusual vocalization similar to the two-voiced songs found in some birds.<ref>{{Cite journal|title = Animal communication: Complex call production in the túngara frog|journal = Nature|date = 2006-05-04|issn = 0028-0836|pages = 38|volume = 441|issue = 7089|doi = 10.1038/441038a|first1 = M.|last1 = Gridi-Papp|first2 = A. S.|last2 = Rand|first3 = M. J.|last3 = Ryan|pmid=16672962|bibcode = 2006Natur.441...38G|s2cid = 4381969|doi-access = free}}</ref>
 
In the [[common toad]], sexual competition is driven in large part by fighting—successful males often physically displaced other males from the backs of a female in order to gain access to mating with that female. Larger males were more successful in such takeovers, and had higher reproductive success as a result.<ref>{{Cite journal|title = Competitive mate searching in male common toads, Bufo bufo|journal = Animal Behaviour|pages = 1253–1267|volume = 27|doi = 10.1016/0003-3472(79)90070-8|first1 = N.B.|last1 = Davies|first2 = T.R.|last2 = Halliday|date = November 1979|s2cid = 54259148}}</ref> However, the vocalizations of these toads provide a reliable signal of body size and thus fighting ability, allowing contests for possession of females to be settled without risk of injury.<ref>{{Cite journal|title = Deep croaks and fighting assessment in toads Bufo bufo|journal = Nature|date = 1978-08-17|pages = 683–685|volume = 274|issue = 5672|doi = 10.1038/274683a0|first1 = N. B.|last1 = Davies|first2 = T. R.|last2 = Halliday|bibcode = 1978Natur.274..683D|s2cid = 4222519}}</ref>
 
In the ''[[Mexican burrowing toad]]'', males produce two types of advertisement calls when attracting females for mating. These are the pre-advertisement and advertisement calls, both of which have a different tonality and purpose. The advertisement call is a single tone with an upward tone, with a duration of about 1.36 seconds. The pre-advertisement call is a single short sound without modulation, and is of higher frequency than the [[Dendropsophus ebraccatus|advertisement calls]].<ref>{{cite journal |last1=Sandoval |first1=L. |title=Sexual size dimorphism and acoustical features of the pre- advertisement and advertisement calls of Rhinophrynus dorsalis Duméril & Bibron, 1841 (Anura: Rhinophrynidae)|journal=Mesoamerican Herpetology |date=2015 |s2cid=208303240 |language=en}}</ref> These signals provide reliable signals to females of the strength and ability of males.
 
In the frog species, [[Bibron's toadlet]], males increase frequency of calls in the presence of other members of the species.
 
=== Insects ===
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[[File:Field cricket Gryllus pennsylvanicus.ogg|thumb|Mating call of field cricket, ''Gryllus pennsylvanicus'']]
 
In the field cricket, ''Gryllus integer'', males rub their wings together to create a rapid trill that produces sound.<ref name="Hedrick, A. V. 2000">{{cite journal | last1 = Hedrick | first1 = A. V. | year = 2000 | title = Crickets with extravagant mating songs compensate for predation risk with extra caution | journal = Proceedings of the Royal Society B: Biological Sciences | volume = 267 | issue = 1444| pages = 671–675 | doi=10.1098/rspb.2000.1054 | pmid=10821611 | pmc=1690585}}</ref> Males individually vary in the durations of their trilling or, what is more sophisticatedly called, bout length. The bout length of each male is heritable and passed on to his future offspring. Also, females prefer to mate with males that have longer bout lengths.<ref>{{cite journal | last1 = Hedrick | first1 = AV | year = 1986 | title = Female preferences for male calling bout duration in a field cricket | journal = Behavioral Ecology and Sociobiology | volume = 19 | issue = 1 | pages = 73–77 | doi=10.1007/bf00303845| bibcode = 1986BEcoS..19...73H | s2cid = 26404984 }}</ref> The end result is that males with longer bout lengths produce more offspring than males with shorter bout lengths.
 
Other factors that influence the formation of these bout lengths include temperature and predation. In field crickets, males prefer warmer sites for mating as shown by an increase in the frequency of their mating calls when they were living in warmer climates.<ref>{{cite journal | last1 = Hedrick | first1 = A. | last2 = Perez | first2 = D. | last3 = Lichti | first3 = N. | last4 = Yew | first4 = J. | year = 2002| title = Temperature preferences of male field crickets (Gryllus integer) alter their mating calls | journal = Journal of Comparative Physiology A| volume = 188| issue = 10| pages = 799–805| doi=10.1007/s00359-002-0368-9| pmid = 12466955 | s2cid = 20635779 }}</ref> Predation also affects the mating calls of field crickets. When in a potentially dangerous environment, males cease calling for longer periods of time when interrupted by a predator cue.<ref name="Hedrick, A. V. 2000"/> This suggests that there is an interplay between intensity of mating call and risk of predation.
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==== Birds ====
The feathers, the beak, the feet, and different tools are all used by different bird species to produce mating calls to attract mates. For example, the snipe uses its feathers to produce a [[drumming (snipe)|"drumming"]] sound to attract mates during a special mating dance. Snipes used specialized tail feathers to create a sound described as a "rattle" or "throbbing" noise.<ref>Bahr, P. H. "On the "bleating" or "drumming" of the snipe (Gallinago coelestis)." Proceedings of the Zoological Society of London. Vol. 77. No. 1. Blackwell Publishing Ltd, 1907</ref> Palm cockatoos use sticks to drum on hollow trees, creating a loud noise to attract the attention of mates.<ref>{{cite journal | last1 = Gray | first1 = Patricia M. |display-authors=etal | year = 1915 | title = Enhanced: The Music of Nature and the Nature of Music | journal = Science Online | volume = 291 | pages = 52–54 }}</ref> [[Bustards]] are large, highly terrestrial birds that stamp their feet during mating displays to attract mates.<ref>{{cite journal | last1 = Alonso | first1 = Juan C. |display-authors=etal | year = 2010 | title = Correlates of male mating success in great bustard leks: the effects of age, weight, and display effort | journal = Behavioral Ecology and Sociobiology | volume = 64 | issue = 10| pages = 1589–1600 | doi=10.1007/s00265-010-0972-6| bibcode = 2010BEcoS..64.1589A | hdl = 10261/76985 | s2cid = 8741416 }}</ref> ''[[Mirafra apiata]]'', commonly known as the Clapper lark, engages in a complex display flight that is characterized by the rattling of the wings.<ref>{{cite journal | last1 = Maclean | first1 = Gordon L | year = 1970 | title = Breeding behaviour of larks in the Kalahari Sandveld | url = http://www.the-eis.com/data/literature/Breeding%20behaviour%20of%20larks%20in%20the%20Kalahari%20sandveld_1970.pdf | journal = Annals of the Natal Museum | volume = 20 | pages = 388–401 | access-date = 2019-01-07 | archive-url = https://web.archive.org/web/20190107232837/http://www.the-eis.com/data/literature/Breeding%20behaviour%20of%20larks%20in%20the%20Kalahari%20sandveld_1970.pdf | archive-date = 2019-01-07 | url-status = dead }}</ref>
 
Many species of birds, such as manakins and hummingbirds, use sonation for mating calls. However, peacocks exhibit a feature of sonation that reveals intrasexual and intersexual properties of this type of mating call.<ref name=":1">{{Cite journal|title = Infrasound in mating displays: a peacock's tale|journal = Animal Behaviour|date = 2015-04-01|pages = 241–250|volume = 102|doi = 10.1016/j.anbehav.2015.01.029|first1 = Angela R.|last1 = Freeman|first2 = James F.|last2 = Hare|s2cid = 53164879}}</ref> Males move their feathers to produce a low-frequency sonation ([[infrasound]]) and sonate more frequently in response to a sonation by other males. This is attributable to a male's desire to advertise its presence above other males looking for mates, suggesting that sonation carries an intrasexual function. In addition, females show increased alertness when hearing the infrasound signals produced by males' wing-shaking, which highlights how the two sexes use sonation to interact with each other.<ref name=":1" />
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==== Fish ====
 
While most bird species use their feathers, tools, or feet to produce sounds and attract mates, many fish species use specialized internal organs to sonate. In [[Gadidae|Gadoid fish]], special muscles attached to the swimbladder assist in the production of knocking or grunting sounds to attract mates.<ref>{{cite journal | last1 = Hawkins | first1 = A. D. | last2 = Just Rasmussen | first2 = Knud | year = 1978 | title = The calls of gadoid fish | journal = Journal of the Marine Biological Association of the United Kingdom | volume = 58 | issue = 4| pages = 891–911 | doi=10.1017/s0025315400056848| bibcode = 1978JMBUK..58..891H | s2cid = 84514769 }}</ref>
 
==== Lepidoptera ====
In many lepidoptera species including the adzuki bean borer ([[Ostrinia scapulalis]]), ultrasonic mating calls are used to attract females and keep them motionless during [[copulation (zoology)|copulation]]. These pulses have an average frequency of 40&nbsp;kHz.<ref name=":2">{{Cite journal|last1=Takanashi|first1=Takuma|last2=Nakano|first2=Ryo|last3=Surlykke|first3=Annemarie|last4=Tatsuta|first4=Haruki|last5=Tabata|first5=Jun|last6=Ishikawa|first6=Yukio|last7=Skals|first7=Niels|date=2010-10-04|title=Variation in Courtship Ultrasounds of Three Ostrinia Moths with Different Sex Pheromones|journal=PLOS ONE|volume=5|issue=10|pages=e13144|doi=10.1371/journal.pone.0013144|pmid=20957230|pmc=2949388|issn=1932-6203|bibcode=2010PLoSO...513144T|doi-access=free}}</ref>
 
== Speciation due to mating call differences ==
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{{reflist}}
 
{{Animal communication}}
[[Category:Mating]]
[[Category:Animal developmental biology]]
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