BIOL 307-B05 Assignment 2: Sexual Selection in birds 1 Jennifer Lowe, 24 February 2014 Research surrounding the causes and effects of sexual selection has come a long way since it was of interest to Charles Darwin1, though all known forms are now encompassed with the current description as successful mating variation2. Birds exhibit some elaborate examples of sexual selection that are often results of male competition and/or female choice. In some species the same male ornamentation is used for both competition and female preference3. Male competition takes many forms and it does not always end at mate acquisition. For example, sperm competition can occur when a female has had multiple mating partners4. Sexual selection involving female choice occurs through several direct and indirect mechanisms5 and can act on many different male adaptations such as courtship displays of song6, plumage ornamentation7, physical displays8, nuptial gifts and/or paternal care9. It has also been shown that a species can exhibit multiple female-preferred traits each related to a different aspect of male fitness10. There are trade-offs associated with plumage ornamentation. Bright plumage can be an indicator of male fitness as it is costly to produce and dull plumage may indicate health conditions11, 12. Also, iridescent plumage has been shown to decrease the hydrophobicity of the feathers and water repulsion is important in temperature regulation and feather cleaning13. Sexual selection for exaggerated characters can be dulled when female choice fluctuates14 or when selection for those characters must compete with selection for alternative characters15. Forms of indirect selection that may result in dramatic male characters include the ‘runaway method’ in which female choice and a male character become genetically correlated which causes positive feedback16 and the ‘parasite hypothesis’ (a form of ‘good genes’ hypothesis) in which females show preference for more ornamented males because they are more parasite-resistant17. The mechanisms of sexual selection have been the focus of much debate as many of the factors involved are difficult to measure (e.g. Female choice)5, 6. BIOL 307-B05 Assignment 2: Sexual Selection in birds 2 Jennifer Lowe, 24 February 2014 Annotated Bibliography 1 Darwin, C. 1874. The descent of man: and selection in relation to sex. A.L. Fowle, N.Y. Print. A look at the evolution of human beings from a ‘lower’ organism and an elaborate comparison between human beings and other animals with focus on thought. A great deal of attention is given to the processes of selection with focus on sexual selection. Though Darwin has brought up sexual selection previously, in this book he gives a more detailed description. The modes described are male-male competition or female choice. 2 Searcy, W.A. & Yasukawa, K. 1995. Polygyny and sexual selection in Red-winged Blackbirds. Princeton University Press, Princeton, N.J. Print. A book that uses original and previous research to show how red-winged blackbird (Agelaius phoeniceus) females can benefit and how the effects sexual selection can become greater by mate-sharing. The book is also a great overview of sexual selection and shows how its definition has required much elaboration since first proposed by Charles Darwin. 3 Griggio, M., Serra, L., Licheri, D., Monti, A. & Pilastro, A. 2007. Armaments and ornaments in the rock sparrow: possible dual utility of a carotenoid-based feather signal. Behav Ecol Sociobiol. 61:423-433. http://link.springer.com.ezproxy.library.uvic.ca/article/10.1007%2Fs00265-006-0270-5 [accessed 22 February 2014] A study with rock sparrows (Petronia petronia) showing that males with larger throat patches are superior at obtaining territory, defending territory and attracting mates. They experimented with real birds and dummy birds showing that males appeared to be threatened by other males with larger patches. To show that patch size was correlated to female choice, the patches of males were experimentally manipulated. 4 Birkhead, T. 1998. Sperm competition in birds. Rev Reprod. 3:123-129 http://www.reproduction-online.org.ezproxy.library.uvic.ca/content/revreprod/3/2/123 [accessed 22 February 2014] A review that looks at the process and outcomes of sperm competition in extra-pair partnerships, where female copulates with males other than her bonded mate, in several types of monogamous birds. In these circumstances, males acquire many adaptations to increase their chances of successful fertilization of the females. 5 Kirkpatrick, M., & Ryan, M.J. 1991. The evolution of mating preferences and the paradox of the lek. Nature. 350:33-38. http://www.nature.com.ezproxy.library.uvic.ca/nature/journal/v350/n6313/abs/350033a0.html [accessed 20 February 2014]. An review of hypotheses of evolution due to female preference. The selection processes of interest are direct and two modes of indirect. Direct selection is the result of female preference on the basis of direct benefits to her survival or reproductive success. Indirect selection can be the result of the runaway process, in which a male trait becomes exaggerated when the trait and the preference of the trait coevolve, or parasite hypothesis, when there is an association between parasite resistance and elaborate male displays. BIOL 307-B05 Assignment 2: Sexual Selection in birds 3 Jennifer Lowe, 24 February 2014 6 Catchpole, C.K. 1987. Bird song, sexual selection and female choice. Trends Ecol Evol. 2(4):94-97. http://www.sciencedirect.com.ezproxy.library.uvic.ca/science/article/pii/0169534787901650# [accessed 19 February 2014] Shows how displays by female aquatic warblers and sedge warblers (Acrocephalus schoenobaenus) in response to recordings of male song can be used to study female choice. Without such indication from the females, it is difficult to determine if the selection process is acting on female choice or other sexual selection processes. 7 Petrie, M., Halliday, T. & Sanders, C. 1991. Peahens prefer peacocks with elaborate trains. Anim. Behav. 41:323-331. http://www.sciencedirect.com.ezproxy.library.uvic.ca/science/article/pii/S0003347205804841# [accessed 24 February 2014] A study that tests the mating preference of female peafowl (Pavo cristatus). The authors conclude that the more embellished the male’s train, the higher his mating success. They point out that male competition does not have a significant factor because a female does not succumb to distractions from other males once she has made her choice. 8 Byers, J., Hebets, E. & Podos, J. 2010. Female mate choice based upon male motor performance. Anim. Behav. 79:771-778 http://www.sciencedirect.com.ezproxy.library.uvic.ca/science/article/pii/S0003347210000308# [accessed 20 February 2014] A review that suggests that physical displays by males are more important than ornamentation in regards to female choice. This study also suggests that evolution of ornamentation was to enhance the physical display rather than functioning as stand-alone adaptations. The reasoning surrounding this assumption is that physical displays are a superior method of determining overall fitness of an animal. 9 Wiggins, D.A. & Morris, R.D. 1986. Criteria for female choice of mates: courtship feeding and paternal care in the common tern. Am. Nat. 128(1):126-129. http://www.jstor.org/stable/2461291 [accessed 24 February 2014] Includes previous studies and original research to show how nuptial gifts in the form of courtship feeding by males can provide the female with much needed nutrients after copulation and can indicate the level of paternal care that the male will perform. The authors own study of the common tern (Sterna hirundo) revealed a large positive correlation between courtship feeding and the care that the males provided their young. 10 Taff, C.C., Steinberger, D., Clark, C., Belinsky, K., Sacks, H., Freeman-Gallant, C.R., Dunn, P.O. & Whittingham, L.A. 2012. Multimodal sexual selection in a warbler: plumage and song are related to different fitness components. Anim. Behav. 84:813-821 http://www.sciencedirect.com.ezproxy.library.uvic.ca/science/article/pii/S0003347212003107 [accessed 22 February 2014] Use previous studies on male plumage combined with their own studies on male song to show that female choice in the socially monogamous common yellowthroats (Geothlypis trichas) acts on both characters. Like many socially monogamous birds common yellowthroats do practice extra-pair mating in which a bonded female will copulate with multiple males. This study found BIOL 307-B05 Assignment 2: Sexual Selection in birds 4 Jennifer Lowe, 24 February 2014 that success in acquiring a bonding mate is related to plumage and success in extra-pair copulation is related to song. 11 Mougeot, F., Perez-Rodriguez, L., Sumozas, N. & Terraube, J. 2007. Parasites, condition, immune responsiveness and carotenoid-based ornamentation in male red-legged partridge Alectoris rufa. J Avian Biol. 40(1):67-74. http://onlinelibrary.wiley.com.ezproxy.library.uvic.ca/doi/10.1111/j.1600048X.2008.04439.x/abstract;jsessionid=E1A84B01902074DC2414B0F7077831D0.f02t03 [accessed 23 February 2014] A study that links health and immunity with ornamentation of carotenoid base. As the researchers predicted, red-legged partridge (Alectoris rufa) males showing higher levels of carotenoids in their ornaments showed a greater immune response. Coccidia parasite infections were used to show the relationship between immune response and carotenoid levels. Another factor that makes brighter ornamentation a good selection factor is that carotenoids must be ingested from food hence revealing how apt a male is at survival. 12 Seifferman, L., Hill, G.E. 2005. Evidence for sexual selection on structural plumage coloration in female eastern bluebirds (Sialia sialis). Evol. 59(8):1819-1828. http://www.jstor.org.ezproxy.library.uvic.ca/stable/3449084 [accessed 23 February 2014] A study showing that female fitness of eastern bluebirds (Sialia sialis) can be indicated by the quality of melanin-based plumage. This study is interesting in that it shows sexual selection acting on female characters as well as those of males. Laboratory experiments determined the impact of nutritional stresses on plumage colouration. Field data was collected to evaluate the relationships between plumage colour, mate choice and mating success. 13 Eliason, C. & Shawkey, D. 2011. Decreased hydrophobicity if iridescent feathers: a potential cost of shiny plumage. J Exp Bio. 214:2157-2163. http://jeb.biologists.org.ezproxy.library.uvic.ca/content/214/13/2157 [accessed 23 February 2014] A study testing the hydrophobicity and self-cleaning efficiency of iridescent feathers and noniridescent feathers. Iridescent feathers have increased surface area due to films of keratin and melanin coating the barbules resulting in the absorption of water. This is considered a cost because birds rely on their feather to repel water which is a temperature-regulating mechanism and self-cleaning mechanism. 14 Caine, A.S. & Lyon, B. 2008. Adaptive plasticity in female mate choice dampens sexual selection on male ornaments in the lark bunting. Science. 319(5862):459-462 http://www.jstor.org/stable/20053208 A study of the lark bunting (Calamospiza melanocorys) displaying that female preference for male plumage characters can change from year to year and the result of this preference variation is that sexual selection for those characters can be dulled. The difference in selection for these traits can be so extreme that a trait can reverse from positive to negative selection from one year to the next which may result in males to develop multiple characters that are sexually selected for. In a dynamic environment, socially or ecologically, it is beneficial for the female to have the ability to choose a male with characters that meet her changing needs. BIOL 307-B05 Assignment 2: Sexual Selection in birds 5 Jennifer Lowe, 24 February 2014 15 Greene, E., Lyon, B.E., Muehter, V.R., Ratcliffe, L., Oliver, S.J. & Boag, P.T. 2000. Disruptive sexual selection for plumage coloration in a passerine bird. Nature. 407(6807):1000http://www.nature.com.ezproxy.library.uvic.ca/nature/journal/v407/n6807/full/4071000a0.html [accessed 18 February 2014] A study that shows how different sexual selection process can result in disruptive selection where both extremes of a character are selected for. Lazuli buntings (Passerina amoena) display sexual selection for male competition and female choice. Young males with plain markings are able to acquire favourable territory because other males do not see them as a threat, whereas bright males are more likely to be favoured by females. The result is a bimodal distribution in plumage coloration. 16 Hall, D.W., Kirkpatrick, M. & West, B. 2000. Runaway sexual selection when female preferences are directly selected. Evolution. 54(6):1862-1869. http://www.jstor.org/stable/2640532 [accessed 20 February 2014] An analysis of runaway selection models where there is coevolution between female preference of a male trait and the trait itself. This mechanism is thought to be responsible for at least some of the most extreme male traits. Runaway can only occur if natural selection does not act negatively on that trait and a stable equilibrium is not reached. This paper looks at many sexual selection scenarios and hypothesizes as to which circumstances are more likely to result in runaway evolution. 17 Hamilton, W.D. & Zuk, M. 1982. Heritable true fitness and bright birds: a role for parasites? Science. 218(4570):384-387. http://www.jstor.org/stable/1688879 [accessed 22 February 2014] The introduction of how female choice for elaborate male plumage may be correlated to male parasite load and genetic resistance to parasites. Their hypothesis includes two predictions; that parasites and their hosts co-evolve providing host with resistance and that when a female chooses her mate on the bases of his showiness, she is also choosing him on the basis of his health (eg. resistance to parasites). BIOL 307-B05 Assignment 2: Sexual Selection in birds 6 Jennifer Lowe, 24 February 2014