User:Alison1 6/sandbox

Final Article Draft

Male display

In most species, the male is the sex that initiates courtship displays in pre-copulatory sexual selection. Performing a display allows the male to present his traits or abilities to a female. Mate choice, in this context, is driven by females. Direct or indirect benefits are often key deciding factors in determining which of these males acquire the opportunity to reproduce and which do not.

Direct benefits can be seen due to the expression of preference. Females can raise their own fitness if they prefer to respond to particular types of signals, independent of costs and certain benefits associated with mating. For example, choosing to mate with males that produce more localized signals would incur less of an energetic investment for a female as she searches for a mate. On the other hand, females can put in more energy towards this process and still attain a higher fitness if they mate with only particular types of males. With this, the males being chosen may impose lower costs on the female or even provide more in terms of material or offspring contributions.

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 an offspring that is potentially more fit.

The male Six-Plumed bird-of-paradise, Parotia lawesii, exemplifies this idea of male courtship display with its ritualized "ballerina dance" and unique occipital and breast feathers that serve to stimulate the female visual system. This stimulation, along with many other factors, results in subsequent copulation or rejection.

contribution

In other species, males may exhibit courtship displays that serve as both visual and auditory stimulation. For example, the male Anna's hummingbird (Calypte anna) and Calliope hummingbird (Stellula calliope) 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 gorget and hovers in front of the female, moving from side to side while rotating his body and tail. The rhythmetic movements of the male's wings produce a distinctive buzzing sound . When conducting a dive display, the male typically ascends approximately 20-35 m in the air then abruptly 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 produces a short, buzzing sound, created by the fluttering and collision of tail feathers.

In addition, some animals attempt to attract females through the construction and decoration of unique structures. For example, male satin bowerbirds (Ptilonorhynchus violaceus) build "bowers" and decorate the structures with colourful and bright objects to attract and stimulate visiting females. Typically, males who acquire the largest number of decorations tend to have greater success in mating.

end

In some species, males initiate courtship rituals only after mounting the female. Courtship may even continue after copulation has been completed. In this systems, the ability of the female to choose their mates is limited. This process, known as copulatory courtship, is prevalent in many insect species.

Extended courtship display [edit]

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 from 15 minutes to a few days. However, certain animals may undergo an extended courtship period, lasting as long as 2 months.

One such exception is the emperor penguin (Aptenodytes forsteri). The emperor penguin engage in an extended courtship period which can last up to 2 months, the longest of any other arctic seabirds. Their courtship period accounts for 16% of the total time they spend breeding, whereas in their closest relatives, the king penguin, the courtship period takes up just 3% of their breeding cycle.

Contribution

Energetic costs of courtship display

Courtship displays typically involve some sort of metabolic cost to the animal performing the display. The energy expended by the animal to perform the courtship behaviour can vary among species. Some animals engage in displays that expend little energy as seen in the salamander (Desmognathus ochrophaeus). Under laboratory settings, courtship behaviours in this species, although complex and involving the release of pheromones, represent as little as approximately 1% of it's daily calorie intake.

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 afterwards. An example of this can be seen in the Greater Sage-Grouse (Centrocercus urophasianus). During the peak of their breeding season, which lasts up to 3 months during spring, leks are frequently visited by groups of up to 70 females. In response to such a large presence of females, males engage in a strutting display up to 6-10 times per minute and continue to perform for approximately 3-4 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.

Mutual display

Often, males and females will perform synchronized or responsive courtship displays in a mutual fashion. With many socially monogamous species such as birds, this duet facilitates pre-copulatory reassurance of pair bonding and strengthens post-copulatory dedication to the development of offspring (e.g. great crested grebe, Podiceps cristatus). For example, male and female crested auklets, Aethia cristatella, will cackle at one another as a vocal form of mutual display which 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 long-tailed manakins, Chiroxiphia linearis.

Wild turkeys (Melagris gallopavo) 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 selection suggests that subordinate males receive indirect benefits by helping related males copulate successfully.

Sexual Ornaments

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 to male supranormal song production.

end

Article Draft

Selected topic: Courtship display.

Sections to add/contribute:

Extended courtship periods

There are several sections within this article that I feel I can contribute, specifically the section discussing "Extended courtship periods." Within this section I plan to add more examples of endurance courtship behaviours and discuss the costs and benefits of such behaviours (e.g., the hummingbird).

Cooperative courtship behaviours

I plan to elaborate on the information regarding cooperative courtship behaviour; providing a more detailed example of the long-tailed manakin, as well as discussing cooperative courtship within the wild turkey species.

Lekking behaviour

I would also like to add some information concerning lekking behaviours observed in birds as it has yet to be touched on ( e.g., use of bowl systems by the New Zealand Kakapo parrot to perform courtship calls and postures).

Female and Male ornaments

Finally, as suggested on this articles talk page, I will add a section highlighting female and male ornaments. (e.g., female mate selection in canaries).

Bibliography

Barske, J., Schlinger, B.A., Wikelski, M., & Fusani, L. (2011) Female choice for male motor skills. Proceedings of the Royal Society B, 278, 3523-3528

Beehler, B. (1983). Lek behavior of the lesser bird of paradise. Auk, 100(4), 992-995.

Bennett, A. F. & Houck, L. D. 1983. The energetic cost of courtship and aggression in a plethodontid salamander. Ecology, 64, 979e983.

Borgia G (1985) Bower quality, number of decorations and mating success of male satin bowerbirds (Ptilonorhynchus violaceus) –an experimental analysis. Anim Behav 33:266–271

Clark, C. (2012). The role of power versus energy in courtship: What is the ‘energetic cost’ of a courtship display? Animal Behaviour, 84(1), 269-277.

Clark, C. 2011. Wing, tail, and vocal contributions to the complex signals of a courting Calliope Hummingbird. Current Zoology, 57: 187-196.

Clark, C., Feo, T., & Escalante, I. (2011). Courtship displays and natural history of scintillant (selasphorus scintilla) and volcano (s. flammula) hummingbirds. The Wilson Journal of Ornithology, 123(2), 218-228.

Garcia-Fernandez, Draganoiu, Ung, Lacroix, Malacarne, & Leboucher. (2013). Female canaries invest more in response to an exaggerated male trait. Animal Behaviour,85(3), 679-684.

Gibson, R. M., J. W. Bradbury and S. L. Vehrencamp. 1991b. Mate choice in lekking Sage Grouse revisited: the roles of vocal display, female site fidelity, and copying. Behav. Ecol. no. 2:165-180.

Hamilton, W. D. The genetical theory of social behavior. I and II. J. Theor. Biol. 7, 1–52 (1964)

Houck, L., Arnold, S., & Thisted, R. (1985). A Statistical Study of Mate Choice: Sexual Selection in a Plethodontid Salamander (Desmognathus ochrophaeus). Evolution, 39(2), 370-386. doi:10.2307/2408370

Lukianchuk, K. C., & Doucet, S. M. (2014). Cooperative courtship display in Long-tailed Manakins Chiroxiphia linearis: Predictors of courtship success revealed through full characterization of display. Journal of Ornithology, 155(3), 729-743.

McDonald, D. B. & Potts, W. K. Cooperative display and relatedness among males in a lek-mating bird. Science 266, 1030–1032 (1994).

Madden, J. (2003). Male spotted bowerbirds preferentially choose, arrange and proffer objects that are good predictors of mating success. Behavioral Ecology and Sociobiology, 53(5), 263-268.

Merton, D., Morris, R., & Atkinson, I. (1984). Lek behaviour in a parrot:the Kakapo Strigops habroptilus of New Zealand. Ibis, 126(3), 277-283.

Krakauer, A. H. (2005). Kin selection and cooperative courtship in wild turkeys. Nature, 434(7029), 69-72.

Scott, J. (1942). Mating Behavior of the Sage Grouse. The Auk, 59(4), 477-498.

Tibbetts, E., Forrest, T., Vernier, C., Jinn, J., & Madagame, A. (2015). Socially selected ornaments and fitness: Signals of fighting ability in paper wasps are positively associated with survival, reproductive success, and rank. Evolution; International Journal of Organic Evolution, 69(11), 2917-26.

Vehrencamp, S.L., Bradbury, J.W., & Gibson, R.M. (1989). The energetic cost of display in male sage grouse. Animal Behaviour, 38, 885-8964

Watts, C. R. & Stokes, A. W. The social order of turkeys. Sci. Am. 224, 112–118 (1971)

Wheeler, J., Gwynne, D., & Bussiere, L. (2012). Stabilizing sexual selection for female ornaments in a dance fly. Journal of Evolutionary Biology, 25(7), 1233-1242.

Zuk, M., Popma, S., & Johnsen, T. (1995). Male courtship displays, ornaments and female mate choice in captive red jungle fowl. Behaviour, 132, 821-836.

Young, J. R., J. W. Hupp, J. W. Bradbury and C. E. Braun. 1994a. Phenotypic divergence of secondary sexual traits among Sage Grouse, Centrocercus urophasianus, populations. Anim. Behav. no. 47:1353-1362.

Article Evaluation

The Article I have selected to evaluate is Courtship display.

• Is everything in the article relevant to the article topic?
  • It appears that all of the material in this article is relevant to the topic. I think that the section on evolutionary significance could be elaborated on as it only focuses on two hypothesis.
• Is there anything that distracted you?
  • Not particularly.
• Is the article neutral? Are there any claims, or frames, that appear heavily biased toward a particular position?
  • The article does a great job providing neutral unbiased information.
• Are there viewpoints that are overrepresented, or underrepresented?
  • The section on multi-modal signal processing is fairly detailed, providing several examples. I feel that the sections discussing evolutionary significance and environmental factors could include more information and examples.
• Check a few citations. Do the links work? Does the source support the claims in the article?
  • Of the citations I checked, the sources supported the information provided in the article. However, not all of the citations had appropriate doi links.
• Is each fact referenced with an appropriate, reliable reference? Where does the information come from? Are these neutral sources? If biased, is that bias noted?
  • No, the author provides several facts throughout the article without proper citations. An example of this is evident in the section discussing male display; "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 an offspring that is potentially more fit." The author does not provided a reference which leads me to question where the information came from?
• Is any information out of date? Is anything missing that could be added?
  • The references provided show articles dating from 2015 as far back as 1970. I think the addition of some more recent sources could benefit this article.
• Check out the Talk page of the article. What kinds of conversations, if any, are going on behind the scenes about how to represent this topic?
  • There are several comments on this articles talk page suggestion the addition of more information such as adding information pertaining to group strategies used by animals to increase their chances of reproductive success. Another comment suggests creating a section for courtship in animals as the article focuses mainly on birds.
• How is the article rated? Is it a part of any WikiProjects?
  • This article is part of the WikiProject Birds. It is rated as start-class on the project's quality scale and rated as low importance on the project's importance scale.
• How does the way Wikipedia discusses this topic differ from the way we've talked about it in class?
  • The information pertaining to multi-modal signal processing has yet to be covered in class.

Article Evaluation (2)

The proposed additions fit well with the main article. There's no bias, just relevant examples illustrating the article's main topic. The talk page does mention that the article is mainly about birds/vertebrates, so maybe including an invertebrate example would help balance the skew in the article. The citations support the statements in the outline. The draft sounds good so far.

Response to feedback

Thank you for your feedback. The suggestion to include an invertebrate example is great, and I will definitely consider including it in my final draft.