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Structural Plumage Colour As a Signal of Mate Quality in Tree Swallows

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Structural Plumage Colour As a Signal of Mate Quality in Tree Swallows 1 Structural Plumage Colour as a Signal of Mate Quality in Tree Swallows Roslyn Dakin A thesis submitted to the Department of Biology in partial fulfillment of the requirements for the degree of Bachelor of Science (Honours) Queen’s University Kingston, Ontario, Canada April 2006 2 ABSTRACT Ornamental plumage colour has long been implicated in sexual selection, and is now thought to function as an honest signal of individual quality in a number of bird species. The purpose of this thesis was to determine whether the bright blue-green structural plumage of tree swallows (Tachycineta bicolor) might function as a signal of mate quality in both males and females. Reflectance spectrometry was used to quantify the plumage colour of breeding adult tree swallows in an Ontario population. Results are discussed with respect to mate choice and sexual selection for honest signaling. First, adult male and female tree swallows differ in plumage colour, with males tending to display bluer, more saturated colour. This sexual dichromatism suggests that structural plumage colour may be sexually selected in tree swallows. Second, birds were found to mate assortatively with respect to plumage colour of the rump region. Tree swallows may therefore use the assessment of plumage colour in mate choice, whether this choice is mutual or unidirectional. Third, relationships were found between plumage colour and body size in males, between plumage colour and ectoparasite load in females, and between reproductive performance and plumage colour in both sexes. Structural plumage colour may therefore be an honest indicator of mate quality in both male and female tree swallows. Tree swallows may benefit by choosing bluer, more intensely-coloured mates in terms of direct or indirect benefits for their offspring. 3 ACKNOWLEDGEMENTS I thank Bob Montgomerie, my supervisor, for providing me with the amazing opportunity to work on this project and others, for providing guidance and support along the way, and for trusting me to drive the biology van throughout the spring. I am also grateful to Dr. Steve Lougheed and Briar Howes for teaching me a great deal about field biology and for supporting my failed attempts to catch skinks. Thanks also to Dr. Vicki Friesen for your input, encouragement and of course, your time in agreeing to be my committee member. I am very grateful to Dr. Mary Stapleton for taking me along during her work on the tree swallows at QUBS; thank you Mary for teaching me so much about these birds, and for sharing your data. Without your guidance this project would have been impossible! Thank you also to Jason Clarke for your field assistance; I appreciated both your expertise with the spectrometer and your great company. I am grateful to the rest of the Montgomerie lab, including Christina Cliffe, Kim MacDonald, Katrina Stockely, Ann McKellar, Kamini Persaud, and Nicole Mideo. Thanks for helping with field and lab work, answering questions, and generally keeping me entertained throughout my time in the lab. Thanks to my wonderful friends for their support and valuable suggestions for my seminar and poster. And finally, I am forever grateful to my parents who suffered through editing this thesis and most of the other things I have had to write throughout my four years at Queen’s. Even when you fell asleep while reading my essays, I was glad to have your help. 4 TABLE OF CONTENTS Abstract ……...…………………………………………………………………….. 2 Acknowledgements ….…………………………………………………………….. 3 List of Figures …………….……………………………......................................... 5 List of Tables ……….…….……………………………………………………….. 6 Introduction and Literature Review ...……………………………………………... 7 Methods …………………………………………………………………………… 18 Results …………………………………………………………………………….. 22 Discussion …………………………………………………………………………. 27 Literature Cited ……………………………………………………………………. 36 Summary …………………………………………………………………………... 41 Appendix …………………………………………………………………………... 53 5 LIST OF FIGURES Figure 1. Spectral ranges of human and avian colour vision ……………………… 42 Figure 2. Representative reflectance spectra for adult male and female tree swallows …………………………………………………………………... 43 Figure 3. Relationship between the number of feather mites of males and females in mated pairs of tree swallows …………….……………………………... 44 Figure 4. Relationships between plumage colour variables of males and females in mated pairs of tree swallows ….…………………………………… ……... 45 Figure 5. Relationships between plumage colour and morphological variables in male tree swallows …...…………………………………………………. 46 Figure 6. Relationships between reproductive performance and plumage colour in both males and females ……...…………………………………………. 47 6 LIST OF TABLES Table 1. Plumage colour variables in adult male and female tree swallows ……… 48 Table 2. Tests of assortative mating by morphological variables in adult tree swallow pairs ……………………………………………………………… 49 Table 3. Tests of assortative mating by plumage colour variables in adult tree swallow pairs ……………………………………………………………… 50 Table 4. Relationships between plumage colour and morphological variables in male and female tree swallows ……………………………………………. 51 Table 5. Relationships between reproductive performance and plumage colour in male and female tree swallows …………………………………………. 52 7 INTRODUCTION AND LITERATURE REVIEW Sexual Selection and Ornamental Plumage The study of sexual selection has long been associated with questions regarding the evolution of ornamental plumage. Darwin pointed to numerous cases of sexual dimorphism in bird plumage, and noticed that often the male bears more conspicuously ornamented plumage than his mate (Darwin, 1871). He recognized that something other than natural selection must be responsible for maintaining costly traits such as long tails and brightly coloured ornaments. This led Darwin to suggest his theory of sexual selection: ornaments and behaviours that improve the ability to attract or compete for mates are favoured because they improve reproductive success despite costs to survival. More recently, different theories have attempted to explain how ornamental traits could be produced and maintained by sexual selection. Fisher’s theory of runaway sexual selection proposes that an arbitrary ornamental trait in one sex may become exaggerated over time if it becomes associated with the genes for the preference of the trait in the other sex (Fisher, 1958). Under this model, the sole advantage of mating with an ornamented individual is that offspring will be more attractive and will therefore have greater reproductive success. Alternatively, Zahavi proposed that ornamental traits that reduce survival are favoured only if they are honest signals of mate quality (Zahavi, 1975). Under this model, only individuals in the best condition are able to express exaggerated ornaments; the preference for more ornamented mates is adaptive in terms of indirect benefits of genes inherited by the offspring, and possibly direct benefits of increased parental care as well. Hamilton and Zuk (1982) suggested that ornamental plumage may function as an honest signal of disease resistance, since there is evidence that the extent of 8 bright plumage is positively correlated with the degree of infection by blood parasites across species. Sexual selection for ornamental plumage may thus be mediated by the genetic benefits of choosing a mate that is resistant to parasites. Although in many species the male is selected to express a higher degree of ornamentation than the female, plumage characteristics may signal female quality as well. Mutual mate choice by plumage colour is predicted to occur in socially monogamous species where both sexes invest heavily in parental care, because both males and females stand to benefit from the careful selection of mates (Johnstone et al., 1996). For example, experimental work with the blue tit (Parus caeruleus) has shown that both females and males prefer partners that have plumage with greater UV-reflectance (Hunt et al., 1999). In species where such mutual mate choice is beneficial, patterns of assortative mating are predicted. Assortative mating by plumage colour has been observed in blue tits (Andersson et al., 1998) and American goldfinches, Carduelis tristis (MacDougall and Montgomerie, 2003). MacDougall and Montgomerie also suggest that assortative mating by plumage colour could have a role in maintaining the variation in ornamental traits despite strong directional selection. The Measurement of Plumage Colouration Many early studies of plumage ornaments assumed that human perception could effectively describe the quality and variation in plumage colour, either by comparison to colour standards or direct scoring by a human observer (such as Hamilton and Zuk, 1982). However, these methods do not necessarily represent the colour stimulus as it is perceived by birds due to fundamental differences between avian and human visual systems. The 9 human visual system consists of three types of photoreceptors or cones that are maximally sensitive to red, green and blue light respectively. The avian system consists of four types of cones: red, green, blue, and an additional cone sensitive to ultraviolet (UV) wavelengths (Bennett and Cuthill, 1994). As a result, birds are sensitive to a much larger range of wavelengths (320-700 nm) extending into the short-wave, UVA portion of the spectrum, whereas humans are sensitive to a more limited range (400-700 nm; see Figure 1). Moreover, due to this additional cone type, birds are able to perceive an additional dimension of colour not
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