Mating call

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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 females. 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.

Contents

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.

Vocalizations

Birds

Song sparrow Song Sparrow (8381415679).jpg
Song sparrow
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. [1] [2]

Large song repertoires are preferred by females of many avian species. [3] One hypothesis for this is that song repertoire is positively correlated with the size of the brain's song control nucleus (HVC). A large HVC would indicate developmental success. In song sparrows, 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. [3]

Japanese bush warbler songs from island populations have an acoustically simple structure when compared to mainland populations. [4] 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 fairywrens; 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. [5] 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. [1]

Red deer stag during rut Red deer stag 2009 denmark.jpg
Red deer stag during rut

Bird calls are also known to continue after pair formation in several socially monogamous bird species. In one experimental population of zebra finches, there was increased singing activity by the male after breeding. [6] 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 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. [6]

Mammals

During the breeding season, mammals will call out to the opposite sex. Male koalas 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. [7] 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. [8] Acoustic signaling is a type of call that can be used from a significant distance encoding an organism's location, condition and identity. [9] Sac-winged bats 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'. [10] Red deer and spotted hyenas along with other mammals also perform acoustic signaling. [11] [12]

Tungara frog Tungara Frog (Engystomops pustulosus) Calling.jpg
Tungara frog

Amphibians

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 [13]

In the 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. [14]

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. [15] 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. [16]

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 advertisement calls. [17] 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

While mating calls in insects are usually associated with mechanical mating calls, such as in crickets, several species of insects use vocalizations to attract mates. In the Asian corn borer, males emit clicking sounds that mimic the echolocation of bats which prey on the moths. They then take advantage of the female's "freezing" response to mate with the female.

In the Japanese lichen moth, however, the female is able to distinguish between the sounds made by males and those made by bats and other predators. As a result, the males use ultrasonic clicking as a more conventional mating signal, compared to the "deceptive" courtship song used in the Asian Corn Bearer. [18]

Mechanical calls

Mating calls also take form through mechanical processes. Animals that are unable to vocalize their call may use their body to attract mates.

Crickets

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. [19] 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. [20] 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. [21] 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. [19] This suggests that there is an interplay between intensity of mating call and risk of predation.

Sonation

As described in Sonation, "the term sonate is described as the deliberate production of sounds, not from the throat, but rather from structures such as the bill, wings, tail, feet and body feathers, or by the use of tools". In several amphibian and fish species, other special structures are used to produce different sounds to attract mates. Birds are common users of sonation, although several amphibian and fish species have been shown to use sonation as a form of mating call as well. In general, sonation is one factor that plays into how a female may choose a mate. There are other features of mating such as territory defense or mate defense, which contribute to the cause of finding suitable mates.

As outlined below, each species uses a distinct method to produce a non-vocal mating call in order to be most successful in attracting mates. The examples below represent the most common examples found in the literature, although many more examples may exist in nature that are still currently unknown.

Feather of male Pavo cristatus (Indian peafowl). These feathers are used in sonation to create infrasound with intent of mating. Feather of male Pavo cristatus (Indian peafowl).jpg
Feather of male Pavo cristatus (Indian peafowl). These feathers are used in sonation to create infrasound with intent of mating.

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" 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. [22] Palm cockatoos use sticks to drum on hollow trees, creating a loud noise to attract the attention of mates. [23] Bustards are large, highly terrestrial birds that stamp their feet during mating displays to attract mates. [24] Mirafra apiata , commonly known as the Clapper lark, engages in a complex display flight that is characterized by the rattling of the wings. [25]

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. [26] 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. [26]

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 Gadoid fish, special muscles attached to the swimbladder assist in the production of knocking or grunting sounds to attract mates. [27]

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. These pulses have an average frequency of 40 kHz. [28]

Speciation due to mating call differences

Differences in mating calls can lead to the separation of different populations within a species. These differences can be due to several factors, including body size, temperature, and other ecological factors. These can arise in the form of tonal, temporal, or behavioral variations in mating calls that subsequently lead to the separation of populations. The separation of these populations due to differences in mating call and mating call preferences can lead to the evolution and creation of new, unique species.

This type of speciation is most often sympatric speciation: where two or more species are created from an existing parent species that all live in the same geographic location. Although there is an absence of research on mammals and birds, this phenomenon has been heavily researched in several frog species around the world. The examples below illuminate speciation due to mating call differences in several frog species around the world. These distinct species are included because they are the focus of the majority of current research.

Two Microhyla olivacea in a mating position Gastrophryne olivacea01a.jpg
Two Microhyla olivacea in a mating position

Microhyla olivacea and Microhyla carolinensis

These two species of narrow-mouthed frog live in the southern United States and have overlapping ranges in Texas and Oklahoma. Researchers have discovered that these two different species alter the frequencies of their call in the overlap zone of their ranges. For example, the Microhyla olivacea mating call has a significantly lower midpoint frequency in the overlap zone than the mating call outside this zone. This leads researchers to suggest that the differences in mating call in the overlap zone of M. olivacea and M. carolinensis act as an isolating mechanism between the two species. They also hypothesize that the evolution of these differences in mating call led to the separation of these two different frog species from one common species. [29]

Engystomops petersi Engystomops petersi 1.jpg
Engystomops petersi

Engystomops petersi

Female preferences for specific male mating calls can lead to sexual selection in mating calls. Females may prefer a specific type of call that certain males possess, in which only those males will be able to mate with females and pass on their genes and specific mating call. As a result, this female preference may lead to divergence of two species.

In Amazonian frogs, sexual selection for different calls has led to the behavioral isolation and speciation of the túngara frog (Engystomops petersi). [30] From genetic and mating call analysis and, researchers were able to identify that two populations of the túngara frog were almost completely reproductively isolated. From their research, scientists believe that differences in female preferences for mating call type have led to the evolution of this speciation process. Specifically, the Yasuní population females prefer the male mating call that includes a whine, while the other population does not prefer this whine. Subsequently, the Yasuní males include the whine in their call, while the other males do not. For this reason, the differences in call have led to the mechanical separation of this species.

Pseudacris triseriata

A Chorus Frog making a mating call Western Chorus Frog (6922520670).jpg
A Chorus Frog making a mating call

Several studies have shown that the species Pseudacris triseriata (Chorus Frog) can be divided into two subspecies, P. t. maculata and P. t. triseriata, due to speciation events from mating call differences. The Chorus Frog has a very large home range, from New Mexico to Southern Canada. These two subspecies have an overlapping range from South Dakota to Oklahoma. In this overlapping range, both the call duration and the calls per second for each species is very different from outside of this range. This means that calls of these two subspecies are more similar outside of this range, and starkly different within the range. For this reason, scientists suggest that these subspecies evolved from differences in mating call type. [31] Additionally, these subspecies are rarely recorded to have hybrid offspring, which further suggests that there is complete speciation due to mating call differences. The differences in mating calls also help to reinforce the speciation process.

Related Research Articles

<span class="mw-page-title-main">Sexual selection</span> Mode of natural selection involving the choosing of and competition for mates

Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with, and compete with members of the same sex for access to members of the opposite sex. These two forms of selection mean that some individuals have greater reproductive success than others within a population, for example because they are more attractive or prefer more attractive partners to produce offspring. Successful males benefit from frequent mating and monopolizing access to one or more fertile females. Females can maximise the return on the energy they invest in reproduction by selecting and mating with the best males.

<span class="mw-page-title-main">Bird vocalization</span> Sounds birds use to communicate

Bird vocalization includes both bird calls and bird songs. In non-technical use, bird songs are the bird sounds that are melodious to the human ear. In ornithology and birding, songs are distinguished by function from calls.

<span class="mw-page-title-main">Syrinx (bird anatomy)</span> The vocal organ of birds

The syrinx is the vocal organ of birds. Located at the base of a bird's trachea, it produces sounds without the vocal folds of mammals. The sound is produced by vibrations of some or all of the membrana tympaniformis and the pessulus, caused by air flowing through the syrinx. This sets up a self-oscillating system that modulates the airflow creating the sound. The muscles modulate the sound shape by changing the tension of the membranes and the bronchial openings. The syrinx enables some species of birds to mimic human speech.

<span class="mw-page-title-main">American green tree frog</span> Species of amphibian

The American green tree frog is a common arboreal species of New World tree frog belonging to the family Hylidae. This nocturnal insectivore is moderately sized and has a bright green to reddish-brown coloration. Commonly found in the central and southeastern United States, the frog lives in open canopy forests with permanent water sources and abundant vegetation. When defending its territory, the frog either emits aggressive call signals or resolves to grapple with intruders, seldom leading to injury or death. To avoid predation, the frog will leap into the water or jump into the treetops.

<span class="mw-page-title-main">Animal song</span>

Animal song is not a well-defined term in scientific literature, and the use of the more broadly defined term vocalizations is in more common use. Song generally consists of several successive vocal sounds incorporating multiple syllables. Some sources distinguish between simpler vocalizations, termed “calls”, reserving the term “song” for more complex productions. Song-like productions have been identified in several groups of animals, including cetaceans, avians (birds), anurans (frogs), and humans. Social transmission of song has been found in groups including birds and cetaceans.

<span class="mw-page-title-main">Superb lyrebird</span> Species of bird

The superb lyrebird is an Australian passerine songbird, one of two species from the family Menuridae, with the other being the much rarer Albert's lyrebird. It is one of the world's largest songbirds, and is renowned for its elaborate tail and courtship displays, and its excellent mimicry. The species is endemic to Australia and is found in forest in the southeast of the country. According to David Attenborough, the superb lyrebird displays one of the most sophisticated voice skills within the animal kingdom—"the most elaborate, the most complex, and the most beautiful".

<span class="mw-page-title-main">Gray treefrog</span> Species of amphibian

The gray treefrog is a species of small arboreal holarctic tree frog native to much of the eastern United States and southeastern Canada.

<span class="mw-page-title-main">Display (zoology)</span> Set of ritualized behaviours in animals

Display behaviour is a set of ritualized behaviours that enable an animal to communicate to other animals about specific stimuli. Such ritualized behaviours can be visual, but many animals depend on a mixture of visual, audio, tactical and chemical signals. Evolution has tailored these stereotyped behaviours to allow animals to communicate both conspecifically and interspecifically which allows for a broader connection in different niches in an ecosystem. It is connected to sexual selection and survival of the species in various ways. Typically, display behaviour is used for courtship between two animals and to signal to the female that a viable male is ready to mate. In other instances, species may make territorial displays, in order to preserve a foraging or hunting territory for its family or group. A third form is exhibited by tournament species in which males will fight in order to gain the 'right' to breed. Animals from a broad range of evolutionary hierarchies avail of display behaviours - from invertebrates such as the simple jumping spider to the more complex vertebrates like the harbour seal.

<i>Teleogryllus oceanicus</i> Species of cricket

Teleogryllus oceanicus, commonly known as the Australian, Pacific or oceanic field cricket, is a cricket found across Oceania and in coastal Australia from Carnarvon in Western Australia and Rockhampton in north-east Queensland

<i>Anomaloglossus beebei</i> Species of amphibian

Anomaloglossus beebei is a species of frog in the family Aromobatidae. This frog is endemic to Guyana, specifically in the Kaieteur National Park. It mainly survives on the giant bromeliad called Brocchinia micrantha. The phytotelmata of this bromeliad is the site of oviposition and tadpole rearing and are defended over time by the males. The females of this species are more brightly golden coloured whereas males are more of a dull tan with brown pigmentation. Males take care of offspring and are preferred due to the elongation of their calls.

<i>Engystomops petersi</i> Species of amphibian

Engystomops petersi is a species of frog in the family Leptodactylidae. It is found in Amazonian Colombia, Ecuador, and Peru. It is morphologically similar to its sibling species, Engystomops freibergi, and for a period the latter was considered to be a junior synonym of Engystomops petersi. Taxonomy and classification of this species is constantly changing due to the continual evolution of behavioral isolation and rapid speciation in the region. There are also records from the Guianas that have not yet been allocated to either species. Divergence of these two species seems to have been driven by behavioural isolation related to male call characteristics more than geographic isolation.

<span class="mw-page-title-main">Túngara frog</span> Species of amphibian

The Túngara frog is a species of frog in the family Leptodactylidae. It is a small nocturnal terrestrial frog found in Mexico, Central America, and the northeastern regions of South America.

<span class="mw-page-title-main">Courtship display</span> Communication to start a relationship with someone or to get sexual contact

A courtship display is a set of display behaviors in which an animal, usually a male, attempts to attract a mate; the mate exercises choice, so sexual selection acts on the display. These behaviors often include ritualized movement ("dances"), vocalizations, mechanical sound production, or displays of beauty, strength, or agonistic ability.

Frogs and toads produce a rich variety of sounds, calls, and songs during their courtship and mating rituals. The callers, usually males, make stereotyped sounds in order to advertise their location, their mating readiness and their willingness to defend their territory; listeners respond to the calls by return calling, by approach, and by going silent. These responses have been shown to be important for species recognition, mate assessment, and localization. Beginning with the pioneering experiments of Robert Capranica in the 1930s using playback techniques with normal and synthetic calls, behavioral biologists and neurobiologists have teamed up to use frogs and toads as a model system for understanding the auditory function and evolution. It is now considered an important example of the neural basis of animal behavior, because of the simplicity of the sounds, the relative ease with which neurophysiological recordings can be made from the auditory nerve, and the reliability of localization behavior. Acoustic communication is essential for the frog's survival in both territorial defense and in localization and attraction of mates. Sounds from frogs travel through the air, through water, and through the substrate. Frogs and toads largely ignore sounds that are not conspecific calls or those of predators, with only louder noises startling the animals. Even then, unless major vibration is included, they usually do not take any action unless the source has been visually identified. The neural basis of communication and audition gives insights into the science of sound applied to human communication.

Vocal learning is the ability to modify acoustic and syntactic sounds, acquire new sounds via imitation, and produce vocalizations. "Vocalizations" in this case refers only to sounds generated by the vocal organ as opposed to by the lips, teeth, and tongue, which require substantially less motor control. A rare trait, vocal learning is a critical substrate for spoken language and has only been detected in eight animal groups despite the wide array of vocalizing species; these include humans, bats, cetaceans, pinnipeds, elephants, and three distantly related bird groups including songbirds, parrots, and hummingbirds. Vocal learning is distinct from auditory learning, or the ability to form memories of sounds heard, a relatively common trait which is present in all vertebrates tested. For example, dogs can be trained to understand the word "sit" even though the human word is not in its innate auditory repertoire. However, the dog cannot imitate and produce the word "sit" itself as vocal learners can.

<span class="mw-page-title-main">Soundscape ecology</span> Study of the effect of environmental sound on organisms

Soundscape ecology is the study of the acoustic relationships between living organisms, human and other, and their environment, whether the organisms are marine or terrestrial. First appearing in the Handbook for Acoustic Ecology edited by Barry Truax, in 1978, the term has occasionally been used, sometimes interchangeably, with the term acoustic ecology. Soundscape ecologists also study the relationships between the three basic sources of sound that comprise the soundscape: those generated by organisms are referred to as the biophony; those from non-biological natural categories are classified as the geophony, and those produced by humans, the anthropophony.

Female copulatory vocalizations, also called female copulation calls or coital vocalizations, are produced by female primates, including human females, and female non-primates. Copulatory vocalizations usually occur during copulation and are hence related to sexual activity. Vocalizations that occur before intercourse, for the purpose of attracting mates, are known as mating calls.

<span class="mw-page-title-main">Sexual selection in birds</span>

Sexual selection in birds concerns how birds have evolved a variety of mating behaviors, with the peacock tail being perhaps the most famous example of sexual selection and the Fisherian runaway. Commonly occurring sexual dimorphisms such as size and color differences are energetically costly attributes that signal competitive breeding situations. Many types of avian sexual selection have been identified; intersexual selection, also known as female choice; and intrasexual competition, where individuals of the more abundant sex compete with each other for the privilege to mate. Sexually selected traits often evolve to become more pronounced in competitive breeding situations until the trait begins to limit the individual's fitness. Conflicts between an individual fitness and signaling adaptations ensure that sexually selected ornaments such as plumage coloration and courtship behavior are "honest" traits. Signals must be costly to ensure that only good-quality individuals can present these exaggerated sexual ornaments and behaviors.

<span class="mw-page-title-main">Sexual selection in amphibians</span> Choice of and competition for mates

Sexual selection in amphibians involves sexual selection processes in amphibians, including frogs, salamanders and newts. Prolonged breeders, the majority of frog species, have breeding seasons at regular intervals where male-male competition occurs with males arriving at the waters edge first in large number and producing a wide range of vocalizations, with variations in depth of calls the speed of calls and other complex behaviours to attract mates. The fittest males will have the deepest croaks and the best territories, with females making their mate choices at least partly based on the males depth of croaking. This has led to sexual dimorphism, with females being larger than males in 90% of species, males in 10% and males fighting for groups of females.

Condition-dependent signaling is a component of sexual selection. Its exact definition remains elusive due to the vagueness of the terms “condition” and “signaling”. It examines which conditions favor the extravagant displays and ornaments, or signals, some organisms poses, despite their lack of evolutionary value in terms of fitness.

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  19. 1 2 Hedrick, A. V. (2000). "Crickets with extravagant mating songs compensate for predation risk with extra caution". Proceedings of the Royal Society B: Biological Sciences. 267 (1444): 671–675. doi:10.1098/rspb.2000.1054. PMC   1690585 . PMID   10821611.
  20. ↑ Hedrick, AV (1986). "Female preferences for male calling bout duration in a field cricket". Behavioral Ecology and Sociobiology. 19: 73–77. doi:10.1007/bf00303845. S2CID   26404984.
  21. ↑ Hedrick, A.; Perez, D.; Lichti, N.; Yew, J. (2002). "Temperature preferences of male field crickets (Gryllus integer) alter their mating calls". Journal of Comparative Physiology A. 188 (10): 799–805. doi:10.1007/s00359-002-0368-9. PMID   12466955. S2CID   20635779.
  22. ↑ 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
  23. ↑ Gray, Patricia M.; et al. (1915). "Enhanced: The Music of Nature and the Nature of Music". Science Online. 291: 52–54.
  24. ↑ Alonso, Juan C.; et al. (2010). "Correlates of male mating success in great bustard leks: the effects of age, weight, and display effort". Behavioral Ecology and Sociobiology. 64 (10): 1589–1600. doi:10.1007/s00265-010-0972-6. hdl:10261/76985. S2CID   8741416.
  25. ↑ Maclean, Gordon L (1970). "Breeding behaviour of larks in the Kalahari Sandveld" (PDF). Annals of the Natal Museum. 20: 388–401. Archived from the original (PDF) on 2019-01-07. Retrieved 2019-01-07.
  26. 1 2 Freeman, Angela R.; Hare, James F. (2015-04-01). "Infrasound in mating displays: a peacock's tale". Animal Behaviour. 102: 241–250. doi:10.1016/j.anbehav.2015.01.029. S2CID   53164879.
  27. ↑ Hawkins, A. D.; Just Rasmussen, Knud (1978). "The calls of gadoid fish". Journal of the Marine Biological Association of the United Kingdom. 58 (4): 891–911. doi:10.1017/s0025315400056848. S2CID   84514769.
  28. ↑ Takanashi, Takuma; Nakano, Ryo; Surlykke, Annemarie; Tatsuta, Haruki; Tabata, Jun; Ishikawa, Yukio; Skals, Niels (2010-10-04). "Variation in Courtship Ultrasounds of Three Ostrinia Moths with Different Sex Pheromones". PLOS ONE. 5 (10): e13144. Bibcode:2010PLoSO...513144T. doi: 10.1371/journal.pone.0013144 . ISSN   1932-6203. PMC   2949388 . PMID   20957230.
  29. ↑ Blair, Frank (1958). "Mating Call in the Speciation of Anuran Amphibians". The American Naturalist. 92 (862): 27–51. doi:10.1086/282007. S2CID   85030912.
  30. ↑ Boul, Kathryn E.; Funk, W. Chris; Darst, Catherine R.; Cannatella, David C.; Ryan, Michael J. (2007-02-07). "Sexual selection drives speciation in an Amazonian frog". Proceedings of the Royal Society of London B: Biological Sciences. 274 (1608): 399–406. doi:10.1098/rspb.2006.3736. ISSN   0962-8452. PMC   1702375 . PMID   17164204.
  31. ↑ Platz, James E. (1989-08-08). "Speciation within the Chorus Frog Pseudacris triseriata: Morphometric and Mating Call Analyses of the Boreal and Western Subspecies". Copeia. 1989 (3): 704–712. doi:10.2307/1445498. JSTOR   1445498.
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