238:, also known as multi-modal signals. There are two leading hypotheses about the adaptive significance of multi-modal signal processing. The multiple message hypothesis states that each signal that a male exhibits will contribute to a possible mate's perception of the male. The redundant signal hypothesis states that the male exhibits multiple signals that portray the same "message" to the female, with each extra signal acting as a fall-back plan for the male should there be a signaling error. The choosy sex may only evaluate one, or a couple, of traits at a given time when interpreting complex signals from the opposite sex. Alternatively, the choosy sex may attempt to process all of the signals at once to facilitate evaluation of the opposite sex.
218:
558:
144:
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540:), resource availability determines when male emperor penguins will be able to return to their breeding grounds to initiate their courtship rituals. The greater the concentration of resources in their feeding ground, the quicker they will be able to restore their body reserves for winter, and the sooner they will be able to return to their breeding grounds. An early return to their breeding grounds comes with an increased likelihood of finding a mate.
85:
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33:
528:), variation in the light environment plays a huge role in their ability to attract mates. Guppy males alter both their 'courtship mode', whether they perform a full courtship display or try to 'engage' in sneak copulations, and distance from females as light intensity changes. Courtship mode also varies with light spectrum and relates to
280:), females have two ornaments — inflatable abdominal sacs and pinnate tibial scales — that they use as courtship displays in mating swarms. Intermediate variations of such female-specific ornaments are sexually selected for by male dance flies in wild populations. These ornaments may also be a signal of high fecundity in females.
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gene or set of genes will be favoured by female choice over time. This would explain why and how such elaborate traits develop within certain species. However, as time goes on and generations pass, the survival advantage associated with one trait may dissipate due to extreme exaggeration to the point that it decreases fitness.
434:), dominant males exhibit agonistic behavior toward female gorillas at very high rates, with the majority of those interactions being courtship-related. Most documented cases of male gorilla aggression toward females is courtship related and is used primarily as a strategy to prevent females from migrating to another male.
163:) of Australia, males of which build and decorate nest-like structures called "bowers". Bowers are decorated with bright and colourful objects (typically blue in colour) to attract and stimulate visiting females. Typically, males who acquire the largest number of decorations tend to have greater success in mating.
586:
proposes that female selection of a mate is dependent on whether or not the male has genes that would increase the quality of the offspring of the female. In some cases, exaggerated male ornamentation may be indicative to a choosing female that a male who is able to place such a large investment in a
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where reproducing comes at an energy cost to males. Such energy costs can include the effort associated in obtaining nuptial gifts for the female or performing long courtship or copulatory behaviors. An added cost from these time and energy investments may come in the form of increased male mortality
177:
Direct benefits may accrue to the female during male courtship displays. Females can raise their own fitness if they respond to courtship behavior that signals benefits to the female rather than the fitness of the male. For example, choosing to mate with males that produce local signals would require
166:
In some species, males initiate courtship rituals only after mounting the female. Courtship may even continue after copulation has been completed. In this system, the ability of the female to choose her mate is limited. This process, known as copulatory courtship, is prevalent in many insect species.
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and hovers in front of the female, moving from side to side while rotating his body and tail. The rhythmic movements of the male's wings produce a distinctive buzzing sound. When conducting a dive display, the male typically ascends approximately 20–35 m (66–115 ft) in the air then abruptly
351:
Sexual ornaments can serve to increase attractiveness and indicate good genes and higher levels of fitness. When exposed to exaggerated male traits, some females may respond by increasing maternal investments. For example, female canaries have been shown to produce larger and denser eggs in response
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Female courtship display is less common in nature as a female would have to invest a lot of energy into both exaggerated traits and in their energetically expensive gametes. However, situations in which males are the sexually selective sex in a species do occur in nature. Male choice in reproduction
578:
As explained by the
Fisherian runaway model, sexually dimorphic males with exaggerated ornamentation may have been sexually selected for in species with female choice. Fitness of these males would increase, resulting in the proliferation of males with such ornamentation over time. This means that a
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Mating is preceded by a courtship/pairing period in many animal mating systems. It is during this period that sexually mature animals select their partners for reproduction. This courtship period, which involves displays to attract a mate by a member of a species, is usually short, lasting anywhere
270:
on male peacock spiders, the reproductive success of an individual relies heavily on a male spider's ability to combine visual and vibratory displays during courtship. The combination of these displays in courtship offers support both to the redundant signal and multiple messages hypotheses for the
181:
Indirect benefits are benefits that may not directly affect the parents' fitness but instead increase the fitness of the offspring. Since the offspring of a female will inherit half of the genetic information from the male counterpart, those traits she saw as attractive will be passed on, producing
497:
In contrast, species that engage in prolonged or elaborate displays expend considerable amounts of energy and run the risk of developing fatigue. To prepare and prevent such a risk, some animals may gain weight before a courtship period, only to lose the weight afterward. An example of this can be
402:
Although rare, agonistic behavior between males and females during courtship displays is seen in nature. Intraspecific agonistic behavior that results in the death of a combatant is rare because of the associated risk of death or injury. However, agonistic behavior that turns dangerous does occur.
506:
During the peak of their breeding season, which lasts up to three months during spring, leks are frequently visited by groups of up to seventy females. In response to such a large presence of females, males engage in a strutting display up to six to ten times per minute for approximately three to
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comes into play. This is competition between sperm to fertilize an egg, which is very competitive as only a single sperm will achieve union. In some insects, the male injects a cocktail of chemicals in seminal fluid together with sperm. The chemicals kill off older sperm from any previous mates,
342:
also engage in co-operative displays in which small groups of males (typically brothers) work together to attract females and deter other competitive males. In many cases, only one male within the group will mate, typically the dominant male. To explain this behaviour, Hamilton's theory of kin
587:
trait somewhat counterintuitive to survival would carry good genes. For example, the costs associated with bright and complex plumage can be high. Only males with good genes are able to support a large investment into the development of such traits, which, in turn displays their high fitness.
252:
amphibians, such as the green tree frog, may use visual cues as well as auditory signals to increase their chances of impressing a mate. When the calls of the tree frogs were held equal, it was determined that females tended to overlook an auditory-only stimulus in favor of males who combined
441:
systems. For example, males will seek to obtain a certain spot or position to perform their courtship display. The best spots are regions of high contention as many males want them for themselves. Because of this direct conflict, agonistic encounters between males are fairly common.
323:
will cackle at one another as a vocal form of mutual display that serves to strengthen a bond between the two. In some cases, males may pair up to perform mutual, cooperative displays in order to increase courtship success and attract females. This phenomenon can be seen with
253:
auditory/visual multi-modal signals. It was seen that female green tree frogs preferred when males coupled the visual display with the auditory communication, concluding that male green tree frogs that are visually accessible can increase their probability of mating success.
209:), females use a temporary ornament, a striped pattern, to both attract males and intimidate rival females. In this case, the female of a species developed a sexually selected signal which serves a dual function of being both attractive to mates and deterring rivals.
379:
up-regulates the female's egg-laying rate, and reduces her desire to re-mate with another male. The cocktail also shortens the female's lifespan, also reducing her likelihood of mating with other males. Also, some females can get rid of the previous male's sperm.
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Tibbetts, Elizabeth A.; Forrest, Taylor; Vernier, Cassondra; Jinn, Judy; Madagame, Andrew (2015-11-01). "Socially selected ornaments and fitness: Signals of fighting ability in paper wasps are positively associated with survival, reproductive success, and rank".
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460:. Emperor penguins engage in an extended courtship period that can last up to two months, the longest of any Arctic seabird. Their courtship period accounts for 16% of the total time they spend breeding, whereas in their closest relatives, the king penguin
485:
Courtship displays typically involve some sort of metabolic cost to the animal performing it. The energy expended to perform courtship behaviour can vary among species. Some animals engage in displays that expend little energy, as seen in the salamander
229:
Many species of animals engage in some type of courtship display to attract a mate, such as dancing, the creation of sounds, and physical displays. However, many species are not limited to only one of these behaviors. The males of a species across many
420:, it is not uncommon for males to employ this claw in battle for a mate. Even though this claw developed as a weapon, it is also closely linked with the crabs' courtship display: it is waved in a certain pattern to attract females for mating.
416:), males have been sexually selected to have one enlarged claw, which can take up anywhere from a third to a half of their total body mass, and one regular claw. Although the enlarged claw is believed to have developed for use in combat for
590:
An alternative is the sensory exploitation hypothesis, which supposes that sexual preferences are the result of preexisting sensory biases, such as that for supernormal stimuli. These could drive the evolution of courtship displays.
532:
risk. On average, male guppies seek out and spend more time in the environment in which their color pattern is the most visible. Males, in the light environment that made them most visible, copulated with the most females.
135:) perform two types of courtship displays involving a combination of visual and vocal display—a stationary shuttle display and dive display. When engaging in the stationary shuttle display, the male displays a flared
241:
The process of multi-modal signaling is believed to help facilitate the courtship process in many species. One such species in which multi-modal signaling is seen to improve mating success is the green tree frog
302:
Often, males and females will perform synchronized or responsive courtship displays in a mutual fashion. With many socially monogamous species such as birds, their duet facilitates pre-copulatory reassurance of
119:
male courtship display is seen through the male's intricate wing scissoring patterns and rapid sidestepping. These stimulations, along with many other factors, result in subsequent copulation or rejection.
493:
Under laboratory settings, courtship behaviours in this species, although complex and involving the release of pheromones, represent as little as approximately one per cent of its daily calorie intake.
549:
mating displays is influenced by the color of the bracket fungus that it mates and courts upon; these flies choose brackets that are lighter, making their displays more visible to the opposite sex.
427:
species, males direct agonistic behavior toward females prior to courtship behaviors. Such behavior can include aggressive vocalizations, displays, and physical aggression. In the western gorilla
374:
This has many consequences. Courtship displays allow the mate performing the selection to have a means on which to base the copulatory decision. If a female chooses more than one male, then
140:
turns and descends in a dive-like fashion. As the male flies over the female, he rotates his body and spreads his tail feathers, which flutter and collide to produce a short, buzzing sound.
3048:
Young, Jessica R.; Hupp, Jerry W.; Bradbury, Jack W.; Braun, Clait E. (1994). "Phenotypic divergence of secondary sexual traits among sage grouse, Centrocercus urophasianus, populations".
274:
Multi-modal signaling is not limited to males. Females in certain species have more than one trait or characteristic that they use in a courtship display to attract mates. In dance flies (
266:
in appearance and signaling behavior. During courtship, male peacock spiders compete using both visual displays and vibratory signals for intersexual communication. Because of the intense
178:
less energy for a female as she searches for a mate. Males may compete by imposing lower mating costs on the female or even providing material or offspring contributions to the female.
382:
After mating has taken place, males perform various actions to prevent females from mating again. What action is performed depends on the animal. In some species, the male produces a
694:
Riede, Tobias; Forstmeier, Wolfgang; Kempenaers, Bart; Goller, Franz (2015). "The functional morphology of male courtship displays in the
Pectoral Sandpiper (Calidris melanotos)".
507:
four hours per day. This frequent and repetitive behaviour can result in energy expenditures of up to 2524 kJ/day compared to the inactive males that typically expend 1218 kJ/day.
170:
In most species, the male sex initiates courtship displays in precopulatory sexual selection. Performing a display allows the male to present his traits or abilities to a female.
2893:
Houck, Lynne D.; Arnold, Stevan J.; Thisted, Ronald A. (1985). "A Statistical Study of Mate Choice: Sexual
Selection in a Plethodontid Salamander (Desmognathus ochrophaeus)".
3170:
Gamble, S. (2003). "Environmental variation and the maintenance of polymorphism: the effect of ambient light spectrum on mating behaviour and sexual selection in guppies".
2473:
Garcia-Fernandez, V.; Draganoiu, T.I.; Ung, D.; Lacroix, A.; Malacarne, G.; Leboucher, G. (2013). "Female canaries invest more in response to an exaggerated male trait".
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Isenmann, P. (1971). "Contribution a' l'e´thologie et a' l'e´cologie du manchot empereur (Aptenodytes forsteri Gray) a' la colonie de Pointe Ge´ologie (Terre Ade´lie)".
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In some species, physical traits that are sexually selected for in male courtship displays may also be used in agonistic behavior between two males for a mate. In
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217:
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Laird, Krispen L.; Clements, Paul; Hunter, Kimberly L.; Taylor, Ryan C. (2016). "Multimodal signaling improves mating success in the green tree frog (
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In addition, some animals attempt to attract females through the construction and decoration of unique structures. This technique can be seen in the
123:
In other species, males may exhibit courtship displays that serve as both visual and auditory stimulation. For example, the male Anna's hummingbird (
1177:
Madden, Joah R. (2003). "Male spotted bowerbirds preferentially choose, arrange and proffer objects that are good predictors of mating success".
367:
Males: their interest is to mate with a large number of completely faithful females, thus spreading their genes widely throughout a population.
390:, the male provides a huge quantity of sperm, enough to last the female's entire life. In some birds and mammals, the male may participate in
1490:
3249:
772:
862:"Courtship and male-male agonistic behaviour of Cosmophasis umbratica Simon, an ornate jumping spider (Araneae: Salticidae) from Singapore"
174:, in this context, is driven by females; direct or indirect benefits to the female often determine which males reproduce and which do not.
2139:
Zubakin, V. A.; Volodin, I. A.; Klenova, A. V.; Zubakina, E. V.; Volodina, E. V.; Lapshina, E. N. (2010). "Behavior of crested auklets (
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519:, resource and light availability, have an effect on the timing and effectiveness of courtship displays in certain species of animals.
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2689:
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1100:
Clark, Christopher J. (2012). "The role of power versus energy in courtship: what is the 'energetic cost' of a courtship display?".
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Bennett, Albert F.; Houck, Lynne D. (1983-10-01). "The
Energetic Cost of Courtship and Aggression in a Plethodontid Salamander".
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from 15 minutes to a few days. However, certain animals may undergo an extended courtship period, lasting as long as two months.
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Females: their interest is to mate with a large number of fit males, thus producing a large quantity of fit and varied offspring.
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Saeki, Yoriko; Kruse, Kipp C.; Switzer, Paul V. (September 2005). "Physiological costs of mate guarding in the
Japanese beetle (
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is the phenomenon in which the interests of males and females in reproduction are not the same: they are often quite different:
1315:"Direct benefits and the evolution of female mating preferences: conceptual problems, potential solutions, and a field cricket"
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3777:
1856:
557:
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Endler, J. A. (2016). "Predation, light intensity and courtship behaviour in
Poecilia reticulata (Pisces: Poeciliidae)".
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Vehrencamp, Sandra L.; Bradbury, Jack W.; Gibson, Robert M. (1989). "The energetic cost of display in male sage grouse".
1999:"Female preference for multi-modal courtship: multiple signals are important for male mating success in peacock spiders"
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fit offspring. In this case, males may compete during courtship by displaying desirable traits to pass on to offspring.
143:
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1808:"A global analysis of aerial displays in passerines revealed an effect of habitat, mating system and migratory traits"
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In many cases, male courtship displays will cause forms of contest competition to develop. This is often seen within
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selection suggests that subordinate males receive indirect benefits by helping related males copulate successfully.
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can arise if males are the sex in a species that are in short supply, for example, if there is a female bias in the
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2706:
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Kahn, A. T. (2014). "Female preferences for timing in a fiddler crab with synchronous courtship waving displays".
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730:
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773:"Calling and display by peacocks (pavo cristatus) at mansion house historic reserve, kawau island, New Zealand"
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474:
3011:"Mate choice in lekking sage grouse revisited: the roles of vocal display, female site fidelity, and copying"
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Schilthuizen, Menno. 2001. Frogs, flies and dandelions: the making of species. Oxford
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Simmons, L.W. (July 1990). "Nuptial feeding in tettigoniids male costs and the rates of fecundity increase".
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Schilthuizen, Menno. 2001. Frogs, flies and dandelions: the making of species. Oxford
University Press p92
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Lehtonen, Jussi; Parker, Geoff A.; Schärer, Lukas (May 2016). "Why anisogamy drives ancestral sex roles".
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Borgia, Gerald (1985). "Bower quality, number of decorations and mating success of male satin bowerbirds (
276:
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Yasui, Y. (1997). "A "Good-Sperm" model can explain the evolution of costly multiple mating by females".
888:"Sparkling feather reflections of a bird-of-paradise explained by finite-difference time-domain modeling"
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991:"Wing, tail, and vocal contributions to the complex acoustic signals of courting Calliope hummingbirds"
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Parra, Jorge E.; Beltrán, Marcela; Zefania, Sama; Dos
Remedios, Natalie; Székely, Tamás (2014-04-01).
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There are multiple hypotheses about how courtship displays may have evolved in animals, including the
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dance" and unique occipital and breast feathers that serve to stimulate the female visual system. In
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In some species, males will perform ritualized movements to attract females. The male six-plumed
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Lukianchuk, K. C.; Doucet, S. M. (2014). "Cooperative courtship display in Long-tailed
Manakins
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Agonistic behavior in courtship displays is not limited to male-male interactions. In many
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Kokko, Hanna; Monaghan, Pat (March 2001). "Predicting the direction of sexual selection".
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2190:: predictors of courtship success revealed through full characterization of display".
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3088:"Male courtship decisions are influenced by light environment and female receptivity"
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466:, the courtship period takes up just three per cent of their breeding cycle.
1614:
1597:
1397:"Experimental assessment of mating opportunities in three shorebird species"
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create complex multi-component signals that have an effect on more than one
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acts on the display. These behaviors often include ritualized movement ("
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in which an animal, usually a male, attempts to attract a mate; the mate
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2297:
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1857:"Complex signal function: developing a framework of testable hypotheses"
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lek males perform a courtship "strutting" display for visiting females.
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136:
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72:, mechanical sound production, or displays of beauty, strength, or
2364:"Stabilizing sexual selection for female ornaments in a dance fly"
2048:"Stabilizing sexual selection for female ornaments in a dance fly"
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Proceedings of the Royal Society of London B: Biological Sciences
729:
Koch, Rebecca E.; Krakauer, Alan H.; Patricelli, Gail L. (2015).
2777:
Ancel, A. (2013). "The long engagement of the emperor penguin".
231:
200:
rates, putting further strain on males attempting to reproduce.
3533:
3408:
3355:. Princeton University Press. pp. 3, 32, 41, 43–44, 155.
2525:
Sperm competition and its evolutionary consequences in insects
813:"Object carrying as socio-sexual display in an aquatic mammal"
412:
2974:
Scott, John W. (1942). "Mating Behavior of the Sage Grouse".
3529:
2362:
Wheeler, J.; Gwynne, D. T.; Bussière, L. F. (2012-07-01).
108:) exemplifies male courtship display with its ritualized "
2554:
Female control: sexual selection by cryptic female choice
2046:
Wheeler, J.; Gwynne, D. T.; Bussière, L. F. (July 2012).
2707:"Sexual coercion and courtship by male western gorillas"
2755:
West, K. (2009). "Animal behaviour: animal courtship".
2096:
Proceedings of the Royal Society B: Biological Sciences
2003:
Proceedings of the Royal Society B: Biological Sciences
1812:
Proceedings of the Royal Society B: Biological Sciences
3352:
A taste for the beautiful: The evolution of attraction
3207:"Why do aptenodytes penguins have high divorce rates?"
1133:
1131:
2969:
2967:
811:
Martin, A. R.; Da Silva, V. M F; Rothery, P. (2008).
2556:. Princeton, New Jersey. Princeton University Press.
2514:. Princeton, New Jersey. Princeton University Press.
2506:
2504:
1688:
1686:
1684:
515:
Various environmental factors, such as temperature,
3699:
3661:
3567:
3522:
3442:
2837:
2835:
151:makes and uses a bower to attract potential mates.
3250:"The Evolutionary Biology of Colonizing Species"
1806:Mikula, P.; Toszogyova, A.; Albrecht, T (2022).
1485:. Cambridge University Press. pp. 141–164.
1095:
1093:
1091:
1598:"An intimidating ornament in a female pipefish"
984:
982:
980:
271:evolution of multi-modal signaling in species.
1745:"Multi-modal courtship in the peacock spider,
1641:"Sound source perception in anuran amphibians"
3545:
3420:
8:
2581:Crudgington, H. & Siva-Jothy, M.T. 200.
2092:"Evolution of displays within the pair bond"
1855:Hebets, Eileen A.; Papaj, Daniel R. (2004).
2684:. Harvard University Press. pp. 3–22.
1273:, species recognition and sexual selection"
3552:
3538:
3530:
3427:
3413:
3405:
454:One such exception is the emperor penguin
3746:Coloration evidence for natural selection
3290:
3222:
3111:
2527:. Biological Reviews vol. 55 pp. 525–567.
2433:
2379:
2115:
2063:
2022:
1965:
1939:"The use of multiple cues in mate choice"
1831:
1782:
1772:
1664:
1623:
1613:
1071:
1006:
921:
911:
836:
670:
660:
2583:Genital damage, kicking and early death
1038:Clark, Christopher James (2009-09-07).
989:Clark, Christopher James (2011-04-01).
860:Lim, Matthew L. M.; Li, Daiqin (2004).
627:
2682:Sexual Coercion in Primates and Humans
3244:
3242:
3081:
3079:
2772:
2770:
2768:
2766:
1738:
1736:
1596:Berglund, A.; Rosenqvist, G. (2008).
352:to male supranormal song production.
7:
3275:"The evolution of sexual preference"
292:Male and female long-billed curlew,
2597:"Handedness in fiddler crab fights"
1861:Behavioral Ecology and Sociobiology
1699:Behavioral Ecology and Sociobiology
1508:Behavioral Ecology and Sociobiology
1479:"Mate quality and mating decisions"
1179:Behavioral Ecology and Sociobiology
2510:Arnqvist, G. & Rowe, L. 2005.
2090:; Price, T. D.; Lande, R. (2013).
1697:), but may not help small males".
1331:10.1016/B978-0-12-380896-7.00006-X
1242:10.1111/j.1469-185X.1991.tb01133.x
561:Male and female Pigeon Guillemot,
25:
1319:Advances in the Study of Behavior
3812:
3811:
3652:
3587:Aristotelian/Distraction display
3192:10.1046/j.1461-0248.2003.00449.x
2381:10.1111/j.1420-9101.2012.02522.x
2065:10.1111/j.1420-9101.2012.02522.x
1583:10.1111/j.1439-0310.2005.01106.x
1464:10.1046/j.1461-0248.2001.00212.x
498:seen in the greater sage-grouse
398:Agonistic behavior and courtship
315:). For example, male and female
2368:Journal of Evolutionary Biology
2052:Journal of Evolutionary Biology
1645:Current Opinion in Neurobiology
1313:Wagner, William E. Jr. (2011).
575:and the good genes hypothesis.
2313:Journal of Theoretical Biology
1937:Candolin, U. (November 2003).
1142:): an experimental analysis".
1:
3778:Frequency-dependent selection
3149:10.1016/S0003-3472(87)80010-6
2950:10.1016/s0003-3472(89)80120-4
2659:10.1016/j.anbehav.2014.09.028
2624:10.1016/j.anbehav.2015.09.012
2585:. Nature vol 407 pp. 855–856.
2487:10.1016/j.anbehav.2013.01.007
1413:10.1016/j.anbehav.2013.12.030
1156:10.1016/s0003-3472(85)80140-8
1114:10.1016/j.anbehav.2012.04.012
213:Multi-modal signal processing
2333:10.1016/0022-5193(64)90038-4
1774:10.1371/journal.pone.0025390
662:10.1371/journal.pone.0025390
394:with other candidate males.
386:after insemination. In some
129:) and calliope hummingbird (
1639:Bee, Mark A. (April 2012).
869:Raffles Bulletin of Zoology
546:Hirtodrosophila mycetophaga
3880:
3395:birdsofparadiseproject.org
2757:Chelsea House Publications
2523:Parker, Geoffrey A. 1970.
1657:10.1016/j.conb.2011.12.014
1280:Journal of Insect Behavior
786:(1): 27–34. Archived from
567:, mutual courtship display
298:, mutual courtship display
221:Bosc's fringe-toed lizard
3806:
3650:
2799:10.1007/s00300-013-1285-9
2726:10.1007/s10329-015-0496-9
2204:10.1007/s10336-014-1059-3
2165:10.1134/S1062359010080066
1958:10.1017/S1464793103006158
1873:10.1007/s00265-004-0865-7
1719:10.1007/s00265-016-2160-9
1265:"Copulatory courtship in
1199:10.1007/s00265-003-0582-7
1140:Ptilonorhynchus violaceus
771:Beauchamp, A. J. (2014).
553:Evolutionary significance
502:Centrocercus urophasianus
463:(Aptenodytes patagonicus)
446:Extended courtship period
223:Acanthodactylus boskianus
161:Ptilonorhynchus violaceus
3707:Anti-predator adaptation
1481:. In Bateson, P. (ed.).
489:Desmognathus ochrophaeus
3316:The American Naturalist
1749:(O.P.-Cambridge, 1874)"
1292:10.1023/a:1007710218609
1008:10.1093/czoolo/57.2.187
913:10.1073/pnas.1323611111
584:"good genes" hypothesis
573:Fisherian runaway model
3450:Advertising in biology
3349:Ryan, Michael (2018).
3273:Fisher, R. A. (1915).
3104:10.1098/rspb.2016.0861
3062:10.1006/anbe.1994.1183
3027:10.1093/beheco/2.2.165
2680:Muller, M. N. (2009).
2192:Journal of Ornithology
2108:10.1098/rspb.2012.3020
2015:10.1098/rspb.2015.2222
1908:10.1006/anbe.1999.1242
1824:10.1098/rspb.2022.0370
1056:10.1098/rspb.2009.0508
829:10.1098/rsbl.2008.0067
568:
482:
457:(Aptenodytes forsteri)
299:
277:Rhamphomyia longicauda
226:
195:. This could arise in
152:
95:
45:
3798:Underwater camouflage
3577:Aggressive/Wicklerian
3436:Signalling in biology
3385:featherlightphoto.com
2595:Perez, D. M. (2015).
2552:Eberhard, W.G. 1996.
1625:10.1093/beheco/arn114
1615:10.1093/beheco/arn114
1477:Parker, G.A. (1983).
1263:Hoikkala, A. (2000).
947:Drosophila subobscura
560:
543:The effectiveness of
536:In emperor penguins (
511:Environmental factors
477:
291:
220:
193:operational sex ratio
146:
115:Drosophila subobscura
88:Male peacock spider,
87:
35:
3773:Evolutionary ecology
3758:Deception in animals
3752:Dazzled and Deceived
3712:Animal communication
3086:Cole, G. L. (2016).
2188:Chiroxiphia linearis
538:Aptenodytes forsteri
340:Meleagris gallopavo)
330:Chiroxiphia linearis
326:long-tailed manakins
262:) are exceptionally
3619:Emsleyan/Mertensian
3499:Distraction display
3455:Agonistic behaviour
3184:2003EcolL...6..463G
3092:Biological Sciences
2856:1983Ecol...64..979B
2791:2013PoBio..36..573A
2705:Breuer, T. (2016).
2616:2015AnBeh.110...99P
2325:1964JThBi...7....1H
2286:Scientific American
2247:10.1038/nature03325
2239:2005Natur.434...69K
2157:2010BioBu..37..823Z
1765:2011PLoSO...625390G
1711:2016BEcoS..70.1517L
1575:2005Ethol.111..863S
1520:1990BEcoS..27...43S
1456:2001EcolL...4..159K
1191:2003BEcoS..53..263M
1050:(1670): 3047–3052.
951:Journal of Genetics
904:2014PNAS..111.4363W
750:10.1642/auk-14-63.1
708:10.1642/AUK-14-25.1
653:2011PLoSO...625390G
525:Poecilia reticulata
479:Greater sage-grouse
418:territorial defense
392:agonistic behaviors
321:Aethia cristatella,
309:great crested grebe
295:Numenius americanus
94:, courtship display
44:, courting a female
42:Antilope cervicapra
3763:Deimatic behaviour
3504:Handicap principle
3494:Deimatic behaviour
3205:Bried, J. (1999).
3015:Behavioral Ecology
2822:L'Oiseau et la RFO
2141:Aethia cristatella
2102:(1757): 20123020.
2009:(1820): 20152222.
1946:Biological Reviews
1818:(1973): 20220370.
1602:Behavioral Ecology
1528:10.1007/BF00183312
1267:Drosophila birchii
963:10.1007/bf02982781
616:Courtship disorder
569:
483:
313:Podiceps cristatus
300:
264:sexually dimorphic
227:
153:
96:
46:
3844:Signalling theory
3831:
3830:
3825:
3824:
3793:Signalling theory
3768:Mimicry#Evolution
3741:Community ecology
3736:Animal coloration
3582:Ant/Myrmecomorphy
3487:Courtship display
2426:10.1111/evo.12793
2420:(11): 2917–2926.
1559:Popillia japonica
1492:978-0-521-27207-0
1366:10.1111/evo.12926
898:(12): 4363–4368.
601:Display (zoology)
376:sperm competition
256:Peacock spiders (
206:Syngnathus typhle
132:Stellula calliope
74:agonistic ability
54:display behaviors
50:courtship display
16:(Redirected from
3871:
3859:Animal sexuality
3817:Category mimicry
3815:
3814:
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3160:
3143:(5): 1376–1385.
3137:Animal Behaviour
3132:
3126:
3125:
3115:
3083:
3074:
3073:
3056:(6): 1353–1362.
3050:Animal Behaviour
3045:
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3006:
3000:
2999:
2971:
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2961:
2938:Animal Behaviour
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2647:Animal Behaviour
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2604:Animal Behaviour
2601:
2592:
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2579:
2573:
2563:
2557:
2550:
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2475:Animal Behaviour
2470:
2464:
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2437:
2408:
2402:
2401:
2383:
2374:(7): 1233–1242.
2359:
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2222:
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2145:Biology Bulletin
2136:
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2119:
2084:
2078:
2077:
2067:
2058:(7): 1233–1242.
2043:
2037:
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1994:
1988:
1987:
1969:
1943:
1934:
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1896:Animal Behaviour
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1401:Animal Behaviour
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1349:
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1144:Animal Behaviour
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1102:Animal Behaviour
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347:Sexual ornaments
268:sexual selection
236:sensory modality
100:bird-of-paradise
62:sexual selection
58:exercises choice
21:
3879:
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3400:bbc.co.uk guide
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3347:
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3313:
3312:
3308:
3279:Eugenics Review
3272:
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3267:
3258:
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3248:
3247:
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3224:10.2307/4089382
3204:
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3199:
3172:Ecology Letters
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2988:10.2307/4079460
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2907:10.2307/2408370
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2864:10.2307/1937804
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2512:Sexual conflict
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2278:
2233:(7029): 69–72.
2224:
2223:
2219:
2185:
2184:
2180:
2138:
2137:
2133:
2088:Servedio, M. R.
2086:
2085:
2081:
2045:
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1996:
1995:
1991:
1941:
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1444:Ecology Letters
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1230:Biology Reviews
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995:Current Zoology
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817:Biology Letters
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634:
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629:
624:
597:
564:Cepphus columba
555:
513:
472:
470:Energetic costs
448:
431:Gorilla gorilla
400:
361:Sexual conflict
358:
356:Sexual conflict
349:
317:crested auklets
286:
215:
188:
157:satin bowerbird
149:satin bowerbird
105:Parotia lawesii
82:
28:
23:
22:
15:
12:
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5:
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3700:Related topics
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3639:In vertebrates
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3477:Apparent death
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3374:External links
3372:
3369:
3368:
3361:
3341:
3328:10.1086/286006
3306:
3285:(3): 184–192.
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3217:(2): 504–512.
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3001:
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2963:
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2928:
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2131:
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1989:
1952:(4): 575–595.
1929:
1902:(5): 921–931.
1886:
1867:(3): 197–214.
1847:
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1747:Maratus volans
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823:(3): 243–245.
803:
763:
744:(2): 349–358.
721:
686:
626:
625:
623:
620:
619:
618:
613:
608:
603:
596:
593:
554:
551:
512:
509:
471:
468:
447:
444:
399:
396:
372:
371:
368:
357:
354:
348:
345:
285:
284:Mutual display
282:
259:Maratus volans
214:
211:
197:mating systems
187:
186:Female display
184:
91:Maratus volans
81:
78:
26:
24:
18:Mating display
14:
13:
10:
9:
6:
4:
3:
2:
3876:
3865:
3864:Bird breeding
3862:
3860:
3857:
3855:
3852:
3850:
3847:
3845:
3842:
3841:
3839:
3818:
3809:
3808:
3805:
3799:
3796:
3794:
3791:
3789:
3786:
3784:
3781:
3779:
3776:
3774:
3771:
3769:
3766:
3764:
3761:
3759:
3756:
3754:
3753:
3749:
3747:
3744:
3742:
3739:
3737:
3734:
3732:
3729:
3727:
3723:
3720:
3718:
3715:
3713:
3710:
3708:
3705:
3704:
3702:
3698:
3692:
3689:
3687:
3684:
3682:
3679:
3677:
3674:
3672:
3669:
3668:
3666:
3664:
3660:
3655:
3645:
3642:
3640:
3637:
3635:
3632:
3630:
3627:
3625:
3622:
3620:
3617:
3615:
3612:
3610:
3607:
3603:
3600:
3599:
3598:
3595:
3593:
3590:
3588:
3585:
3583:
3580:
3578:
3575:
3574:
3572:
3570:
3566:
3562:
3555:
3550:
3548:
3543:
3541:
3536:
3535:
3532:
3527:
3525:
3521:
3515:
3512:
3510:
3509:Mobbing calls
3507:
3505:
3502:
3500:
3497:
3495:
3492:
3488:
3485:
3484:
3483:
3480:
3478:
3475:
3471:
3468:
3467:
3466:
3463:
3461:
3458:
3456:
3453:
3451:
3448:
3447:
3445:
3441:
3437:
3430:
3425:
3423:
3418:
3416:
3411:
3410:
3407:
3401:
3398:
3396:
3393:
3391:
3388:
3386:
3383:
3381:
3378:
3377:
3373:
3364:
3362:9780691167268
3358:
3354:
3353:
3345:
3342:
3337:
3333:
3329:
3325:
3321:
3317:
3310:
3307:
3302:
3298:
3293:
3288:
3284:
3280:
3276:
3269:
3266:
3255:
3251:
3245:
3243:
3239:
3234:
3230:
3225:
3220:
3216:
3212:
3208:
3201:
3198:
3193:
3189:
3185:
3181:
3177:
3173:
3166:
3163:
3158:
3154:
3150:
3146:
3142:
3138:
3131:
3128:
3123:
3119:
3114:
3109:
3105:
3101:
3097:
3093:
3089:
3082:
3080:
3076:
3071:
3067:
3063:
3059:
3055:
3051:
3044:
3041:
3036:
3032:
3028:
3024:
3020:
3016:
3012:
3005:
3002:
2997:
2993:
2989:
2985:
2981:
2977:
2970:
2968:
2964:
2959:
2955:
2951:
2947:
2943:
2939:
2932:
2929:
2924:
2920:
2916:
2912:
2908:
2904:
2900:
2896:
2889:
2886:
2881:
2877:
2873:
2869:
2865:
2861:
2857:
2853:
2849:
2845:
2838:
2836:
2832:
2827:
2823:
2816:
2813:
2808:
2804:
2800:
2796:
2792:
2788:
2784:
2780:
2779:Polar Biology
2773:
2771:
2769:
2767:
2763:
2758:
2751:
2748:
2743:
2739:
2735:
2731:
2727:
2723:
2719:
2715:
2708:
2701:
2698:
2693:
2691:9780674033245
2687:
2683:
2676:
2673:
2668:
2664:
2660:
2656:
2652:
2648:
2641:
2638:
2633:
2629:
2625:
2621:
2617:
2613:
2609:
2605:
2598:
2591:
2588:
2584:
2578:
2575:
2572:
2571:0-19-850392-X
2568:
2562:
2559:
2555:
2549:
2546:
2543:
2542:0-19-850392-X
2539:
2533:
2530:
2526:
2520:
2517:
2513:
2507:
2505:
2501:
2496:
2492:
2488:
2484:
2480:
2476:
2469:
2466:
2461:
2457:
2453:
2449:
2445:
2441:
2436:
2431:
2427:
2423:
2419:
2415:
2407:
2404:
2399:
2395:
2391:
2387:
2382:
2377:
2373:
2369:
2365:
2358:
2355:
2350:
2346:
2342:
2338:
2334:
2330:
2326:
2322:
2318:
2314:
2307:
2304:
2299:
2295:
2291:
2287:
2280:
2277:
2272:
2268:
2264:
2260:
2256:
2252:
2248:
2244:
2240:
2236:
2232:
2228:
2221:
2218:
2213:
2209:
2205:
2201:
2197:
2193:
2189:
2182:
2179:
2174:
2170:
2166:
2162:
2158:
2154:
2150:
2146:
2142:
2135:
2132:
2127:
2123:
2118:
2113:
2109:
2105:
2101:
2097:
2093:
2089:
2083:
2080:
2075:
2071:
2066:
2061:
2057:
2053:
2049:
2042:
2039:
2034:
2030:
2025:
2020:
2016:
2012:
2008:
2004:
2000:
1993:
1990:
1985:
1981:
1977:
1973:
1968:
1963:
1959:
1955:
1951:
1947:
1940:
1933:
1930:
1925:
1921:
1917:
1913:
1909:
1905:
1901:
1897:
1890:
1887:
1882:
1878:
1874:
1870:
1866:
1862:
1858:
1851:
1848:
1843:
1839:
1834:
1829:
1825:
1821:
1817:
1813:
1809:
1802:
1799:
1794:
1790:
1785:
1780:
1775:
1770:
1766:
1762:
1759:(9): e25390.
1758:
1754:
1750:
1748:
1739:
1737:
1733:
1728:
1724:
1720:
1716:
1712:
1708:
1704:
1700:
1696:
1689:
1687:
1685:
1681:
1676:
1672:
1667:
1662:
1658:
1654:
1650:
1646:
1642:
1635:
1632:
1626:
1621:
1616:
1611:
1607:
1603:
1599:
1592:
1589:
1584:
1580:
1576:
1572:
1568:
1564:
1560:
1553:
1550:
1545:
1541:
1537:
1533:
1529:
1525:
1521:
1517:
1513:
1509:
1502:
1499:
1494:
1488:
1484:
1480:
1473:
1470:
1465:
1461:
1457:
1453:
1449:
1445:
1438:
1435:
1430:
1426:
1422:
1418:
1414:
1410:
1406:
1402:
1398:
1391:
1388:
1383:
1379:
1375:
1371:
1367:
1363:
1359:
1355:
1348:
1345:
1340:
1336:
1332:
1328:
1324:
1320:
1316:
1309:
1306:
1301:
1297:
1293:
1289:
1285:
1281:
1274:
1272:
1268:
1259:
1256:
1251:
1247:
1243:
1239:
1235:
1231:
1224:
1221:
1216:
1212:
1208:
1204:
1200:
1196:
1192:
1188:
1184:
1180:
1173:
1170:
1165:
1161:
1157:
1153:
1149:
1145:
1141:
1134:
1132:
1128:
1123:
1119:
1115:
1111:
1107:
1103:
1096:
1094:
1092:
1088:
1083:
1079:
1074:
1069:
1065:
1061:
1057:
1053:
1049:
1045:
1041:
1034:
1031:
1026:
1022:
1018:
1014:
1009:
1004:
1000:
996:
992:
985:
983:
981:
977:
972:
968:
964:
960:
956:
952:
948:
941:
938:
933:
929:
924:
919:
914:
909:
905:
901:
897:
893:
889:
882:
879:
875:(2): 435–448.
874:
870:
863:
856:
853:
848:
844:
839:
834:
830:
826:
822:
818:
814:
807:
804:
793:on 2018-02-17
789:
785:
781:
774:
767:
764:
759:
755:
751:
747:
743:
739:
732:
725:
722:
717:
713:
709:
705:
701:
697:
690:
687:
682:
678:
673:
668:
663:
658:
654:
650:
647:(9): e25390.
646:
642:
638:
631:
628:
621:
617:
614:
612:
609:
607:
606:Lek (biology)
604:
602:
599:
598:
594:
592:
588:
585:
580:
576:
574:
566:
565:
559:
552:
550:
548:
547:
541:
539:
534:
531:
527:
526:
520:
518:
510:
508:
505:
503:
495:
492:
490:
480:
476:
469:
467:
465:
464:
459:
458:
452:
445:
443:
440:
435:
433:
432:
426:
421:
419:
415:
414:
409:
408:fiddler crabs
404:
397:
395:
393:
389:
385:
380:
377:
369:
366:
365:
364:
362:
355:
353:
346:
344:
341:
337:
333:
331:
327:
322:
318:
314:
310:
306:
297:
296:
290:
283:
281:
279:
278:
272:
269:
265:
261:
260:
254:
251:
247:
246:
239:
237:
233:
224:
219:
212:
210:
208:
207:
203:In pipefish (
201:
198:
194:
185:
183:
179:
175:
173:
168:
164:
162:
158:
150:
145:
141:
138:
134:
133:
128:
127:
121:
118:
116:
111:
107:
106:
101:
93:
92:
86:
79:
77:
75:
71:
70:vocalizations
67:
63:
59:
55:
51:
43:
39:
34:
30:
19:
3788:Polymorphism
3783:Phagomimicry
3750:
3731:Co-evolution
3486:
3460:Alarm signal
3380:stanford.edu
3351:
3344:
3319:
3315:
3309:
3282:
3278:
3268:
3257:. Retrieved
3253:
3214:
3210:
3200:
3175:
3171:
3165:
3140:
3136:
3130:
3095:
3091:
3053:
3049:
3043:
3018:
3014:
3004:
2979:
2975:
2941:
2937:
2931:
2898:
2894:
2888:
2847:
2843:
2825:
2821:
2815:
2782:
2778:
2756:
2750:
2720:(1): 29–38.
2717:
2713:
2700:
2681:
2675:
2650:
2646:
2640:
2607:
2603:
2590:
2577:
2561:
2548:
2532:
2519:
2478:
2474:
2468:
2417:
2413:
2406:
2371:
2367:
2357:
2316:
2312:
2306:
2289:
2285:
2279:
2230:
2226:
2220:
2195:
2191:
2187:
2181:
2148:
2144:
2140:
2134:
2099:
2095:
2082:
2055:
2051:
2041:
2006:
2002:
1992:
1949:
1945:
1932:
1899:
1895:
1889:
1864:
1860:
1850:
1815:
1811:
1801:
1756:
1752:
1746:
1702:
1698:
1695:Hyla cinerea
1694:
1648:
1644:
1634:
1608:(1): 54–59.
1605:
1601:
1591:
1566:
1562:
1558:
1552:
1514:(1): 43–47.
1511:
1507:
1501:
1482:
1472:
1447:
1443:
1437:
1404:
1400:
1390:
1357:
1353:
1347:
1322:
1318:
1308:
1283:
1279:
1270:
1266:
1258:
1233:
1229:
1223:
1182:
1178:
1172:
1147:
1143:
1139:
1105:
1101:
1047:
1043:
1033:
998:
994:
954:
950:
946:
940:
895:
891:
881:
872:
868:
855:
820:
816:
806:
795:. Retrieved
788:the original
783:
779:
766:
741:
737:
724:
699:
695:
689:
644:
640:
630:
589:
581:
577:
570:
562:
544:
542:
537:
535:
523:
522:In guppies (
521:
514:
499:
496:
487:
484:
462:
456:
453:
449:
436:
428:
422:
411:
405:
401:
381:
373:
359:
350:
339:
336:Wild turkeys
334:
329:
320:
312:
305:pair bonding
301:
293:
275:
273:
257:
255:
245:Hyla cinerea
243:
240:
228:
222:
204:
202:
189:
180:
176:
169:
165:
160:
154:
130:
126:Calypte anna
124:
122:
113:
104:
97:
89:
80:Male display
52:is a set of
49:
47:
41:
29:
3717:Aposematism
3592:Automimicry
3470:Unkenreflex
3465:Aposematism
3443:Non-mimicry
2319:(1): 1–16.
1483:Mate Choice
517:photoperiod
388:hymenoptera
384:mating plug
172:Mate choice
3838:Categories
3722:Camouflage
3691:Vavilovian
3686:Pouyannian
3681:Gilbertian
3644:Wasmannian
3569:In animals
3322:(3): 573.
3259:2019-11-18
2828:: 136–159.
2610:: 99–104.
1561:Newman)".
1271:D. serrata
797:2018-02-13
622:References
439:lek mating
3676:Dodsonian
3663:In plants
3629:Müllerian
3602:Locomotor
3390:bbc.co.uk
3035:1045-2249
2895:Evolution
2872:1939-9170
2653:: 35–39.
2444:1558-5646
2414:Evolution
2390:1420-9101
2255:1476-4687
1421:0003-3472
1407:: 83–90.
1354:Evolution
1064:0962-8452
1017:1674-5507
702:: 65–77.
611:Courtship
530:predation
225:love bite
110:ballerina
38:blackbuck
3854:Ethology
3671:Bakerian
3614:Chemical
3597:Batesian
3514:Stotting
3336:84991216
3301:21259607
3254:epdf.pub
3157:53169408
3122:27683362
3070:54287148
2958:53195569
2923:28564228
2807:15504155
2742:17684689
2734:26483073
2714:Primates
2667:44139315
2632:53194236
2495:53154422
2460:10079068
2452:26456701
2398:22551204
2298:24922757
2263:15744300
2212:15592813
2126:23427172
2074:22551204
2033:26631566
1976:14700392
1967:1975/138
1916:10564594
1881:16494821
1842:35440206
1793:21980440
1753:PLOS ONE
1727:15612885
1675:22265243
1563:Ethology
1544:24088701
1429:15727544
1374:27110661
1339:80868197
1300:25532996
1250:85822041
1236:: 1–31.
1215:44048073
1164:53192968
1122:45711419
1082:19515669
1025:43960957
971:43641513
932:24591592
847:18364306
780:Notornis
758:30509108
716:86155377
681:21980440
641:PLOS ONE
595:See also
248:). Many
3726:Crypsis
3624:Eyespot
3561:Mimicry
3524:Mimicry
3482:Display
3292:2987134
3233:4089382
3211:The Auk
3180:Bibcode
3113:5046890
2996:4079460
2976:The Auk
2915:2408370
2880:1937804
2852:Bibcode
2844:Ecology
2787:Bibcode
2612:Bibcode
2349:5310280
2341:5875341
2321:Bibcode
2271:1457512
2235:Bibcode
2173:9709011
2153:Bibcode
2117:3619481
2024:4685782
1984:8577794
1924:1372407
1833:9019522
1784:3181266
1761:Bibcode
1707:Bibcode
1666:3338885
1571:Bibcode
1536:4600442
1516:Bibcode
1452:Bibcode
1382:2107146
1325:: 273.
1207:4602214
1187:Bibcode
1073:2817121
923:3970480
900:Bibcode
838:2610054
738:The Auk
696:The Auk
672:3181266
649:Bibcode
425:primate
410:(genus
147:A male
3849:Mating
3810:
3634:Sexual
3359:
3334:
3299:
3289:
3231:
3155:
3120:
3110:
3068:
3033:
2994:
2956:
2921:
2913:
2878:
2870:
2805:
2740:
2732:
2688:
2665:
2630:
2569:
2540:
2493:
2458:
2450:
2442:
2396:
2388:
2347:
2339:
2296:
2269:
2261:
2253:
2227:Nature
2210:
2171:
2124:
2114:
2072:
2031:
2021:
1982:
1974:
1922:
1914:
1879:
1840:
1830:
1791:
1781:
1725:
1673:
1663:
1542:
1534:
1489:
1427:
1419:
1380:
1372:
1337:
1298:
1248:
1213:
1205:
1162:
1120:
1080:
1070:
1062:
1023:
1015:
969:
930:
920:
845:
835:
756:
714:
679:
669:
250:anuran
137:gorget
66:dances
3609:Brood
3332:S2CID
3229:JSTOR
3153:S2CID
3066:S2CID
2992:JSTOR
2954:S2CID
2911:JSTOR
2876:JSTOR
2803:S2CID
2738:S2CID
2710:(PDF)
2663:S2CID
2628:S2CID
2600:(PDF)
2491:S2CID
2456:S2CID
2345:S2CID
2294:JSTOR
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