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Causes and Consequences of Blue-Green Eggshell Colour Variation in Mountain Bluebirds (Sialia Currucoides)

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Causes and Consequences of Blue-Green Eggshell Colour Variation in Mountain Bluebirds (Sialia Currucoides) View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by British Columbia's network of post-secondary digital repositories CAUSES AND CONSEQUENCES OF BLUE-GREEN EGGSHELL COLOUR VARIATION IN MOUNTAIN BLUEBIRDS (SIALIA CURRUCOIDES) by Jeannine A. Randall B.Sc., University of Victoria, 2007 THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTERS OF SCIENCE IN NATURAL RESOURCES AND ENVIRONMENTAL STUDIES (BIOLOGY) UNIVERSITY OF NORTHERN BRITISH COLUMBIA August 2016 © Jeannine Randall, 2016 Abstract The function and evolution of ornamental traits has been a major focus of evolutionary ecology. Despite this, female ornaments have received relatively little consideration, and are still poorly understood relative to those produced by males. However, presently, there is much interest in determining how sexual selection shapes female phenotypes. Blue-green eggshell colour, derived from the antioxidant pigment biliverdin, is one attribute produced by female birds that has come under scrutiny as a potentially sexually selected trait. Based on the possibility that biliverdin is limited and costly to produce, the sexually selected egg colour hypothesis predicts that blue-green egg colour has evolved in species with biparental care as an advertisement of female quality that elicits increased paternal effort from their social mate. I combined observations of naturally occurring patterns of eggshell colour variation and parental provisioning rates with an experimental approach to investigate the signalling function of blue-green eggshell colouration in mountain bluebirds (Sialia currucoides). I quantified patterns of within- and among-clutch colour variation and found that eggshell colour was repeatable for individual females, but declined later in the laying sequence within clutches, and between first and second breeding attempts within breeding seasons, suggesting pigments were limited. Using a food and nutrient (carotenoid) supplementation experiment, I determined that eggshell colour is sensitive to food, but not antioxidant availability in this species, and that supplementation did not affect patterns of pigmentation within clutches. Colour of eggshells was not consistently related to female traits such as clutch initiation date and plumage colour, but it did reflect investment in eggs, as eggshell colour was positively related to both egg mass and relative yolk volume. In addition, I conducted a cross-fostering experiment to test whether eggshell colour predicted nestling performance or rearing conditions. I found that eggshell colour was unrelated to nestling ii outcomes, as nestlings hatched from eggs with more saturated blue-green colour did not grow faster and were no more likely to fledge than those hatched from less saturated eggs. I detected relationships between the colour of the eggs in the nest that young were fostered in and their growth, but these associations were not consistent between years. Finally, I used the same cross-fostering design to test the effect of blue-green eggshell colour, separately from the potential influence of nestling phenotypes, on provisioning behaviour of males. I found that the provisioning rates of males during mid brood rearing were not related to the colour of the eggs that young hatched from, but were related to the colour of the eggs that were in their nest during incubation. However, contrary to my predictions if eggshell colour was sexually selected, males fed less at nests with more saturated eggshell colour. Together, my findings do not provide strong support for the sexually selected eggshell colour hypothesis. My results suggest that eggshell pigments are limited and sensitive to food availability and annual variation in conditions, but the inconsistent relationship with nestling performance, and the lack of a positive response by male birds, do not illustrate clear benefits for females to produce blue-green eggs. iii Table of contents Abstract ......................................................................................................................................... ii Table of contents .......................................................................................................................... iv List of tables ................................................................................................................................. vi List of figures ............................................................................................................................. viii Acknowledgements ...................................................................................................................... xi Chapter 1. General introduction ................................................................................................. 1 1.1. Introduction.................................................................................................................. 1 1.2. Study area and species ................................................................................................. 4 1.3. Objectives .................................................................................................................... 5 Chapter 2. Egg colour in mountain bluebirds (Sialia currucoides): patterns of variation and relationships with female quality and investment in eggs ................................................. 8 2.1. Abstract ........................................................................................................................ 8 2.2. Introduction.................................................................................................................. 9 2.3 Methods ...................................................................................................................... 14 2.3.1. Study site, species, and general field procedures ....................................... 14 2.3.2. Colour quantification .................................................................................. 15 2.3.3. Statistical analyse......................................................................................... 16 2.4. Results ....................................................................................................................... 18 2.5. Discussion .................................................................................................................. 29 2.6. Conclusion ................................................................................................................. 32 Chapter 3. An experimental test of the effect of food and antioxidants on blue-green eggshell colouration in mountain bluebirds (Sialia currucoides) ........................................... 34 3.1. Abstract ....................................................................................................................... 34 3.2. Introduction................................................................................................................. 35 3.3. Methods ...................................................................................................................... 37 3.3.1. Study site, species, and general field procedures ....................................... 37 3.3.2. Experimental design ................................................................................... 38 3.3.3. Colour quantification .................................................................................. 39 3.3.4. Statistical analyses ...................................................................................... 40 3.4. Results ....................................................................................................................... 42 3.5. Discussion .................................................................................................................. 47 3.6. Conclusion ................................................................................................................. 50 Chapter 4. Blue-green eggshell colour does not predict nestling performance and may negatively relate to rearing conditions in mountain bluebirds (Sialia currucoides) .............. 52 4.1. Abstract ....................................................................................................................... 52 4.2. Introduction................................................................................................................ 53 4.3. Methods ..................................................................................................................... 56 4.3.1. Study site, species, and general field procedures ....................................... 56 4.3.2. Colour quantification .................................................................................. 58 iv 4.3.3. Experimental design ................................................................................... 59 4.3.4. Statistical analyses ...................................................................................... 59 4.4. Results ....................................................................................................................... 62 4.4.1. Mass growth rate......................................................................................... 62 4.4.2. Primary growth rate .................................................................................... 69 4.4.3. Tarsus growth rate ....................................................................................... 76 4.4.4. Fledging
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