Investigations of Evolutionary Arms
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! INVESTIGATIONS OF EVOLUTIONARY ARMS RACES AND HOST DIVERSITY IN AVIAN BROOD PARASITE SYSTEMS A thesis submitted in partial fulfilment of the requirements for the Degree of Doctor of Philosophy in Biological Sciences at the University of Canterbury by Justin Lee Rasmussen University of Canterbury 2013 ! Table of Contents Acknowledgements........................................................................................................1 Abstract..........................................................................................................................6 Chapter 1: General Introduction ....................................................................................9 Outline of thesis ...................................................................................................................18 Literature cited.....................................................................................................................23 Chapter 2: Does host nest defence behaviour constrain host diversity in avian brood parasites?......................................................................................................................32 2.1 Abstract ..........................................................................................................................32 2.2 Introduction....................................................................................................................33 2.3 Methods..........................................................................................................................36 2.4 Results ............................................................................................................................45 2.5 Discussion ......................................................................................................................46 Literature cited.....................................................................................................................52 Chapter 3: Do red-winged blackbirds Agelaius phoeniceus respond to brown-headed cowbird Molothrus ater eggs with changes in incubation behaviour?. .......................66 3.1 Abstract ..........................................................................................................................66 3.2 Introduction....................................................................................................................67 3.3 Methods..........................................................................................................................71 3.4 Results ............................................................................................................................79 3.5 Discussion ......................................................................................................................81 Literature cited.....................................................................................................................86 Chapter 4: Do shining cuckoos Chrysococcyx lucidus produce preen wax that mimics the preen wax of their host?.........................................................................................95 4.1 Abstract ..........................................................................................................................95 4.2 Introduction....................................................................................................................96 4.3 Methods........................................................................................................................105 4.4 Results ..........................................................................................................................113 4.5 Discussion ....................................................................................................................117 Literature cited...................................................................................................................122 Chapter 5: Does the incubation behaviour of song thrush Turdus philomelos change in response to the odours of conspecifics and heterospecifics?.....................................159 5.1 Abstract ........................................................................................................................159 5.2 Introduction..................................................................................................................160 5.3 Methods........................................................................................................................163 5.4 Results ..........................................................................................................................169 5.5 Discussion ....................................................................................................................170 Literature cited...................................................................................................................173 Chapter 6: General Discussion ..................................................................................182 Literature cited...................................................................................................................194 Appendix 1: Are grey warblers Gerygone igata the optimal host of the shining cuckoo Chrysococcyx lucidus in New Zealand? ....................................................................203 A1.1 Abstract......................................................................................................................203 A1.2 Introduction ...............................................................................................................204 A1.3 Methods .....................................................................................................................205 A1.4 Results........................................................................................................................206 ! A1.5 Discussion..................................................................................................................209 Literature cited...................................................................................................................214 Appendix 2: Do hosts of avian brood parasites respond to artificial eggs and real eggs the same way in artificial parasitism experiments? ...................................................218 A2.1 Abstract......................................................................................................................218 A2.2 Introduction ...............................................................................................................219 A2.3 Methods .....................................................................................................................220 A2.4 Results........................................................................................................................223 A2.5 Discussion..................................................................................................................225 Literature cited...................................................................................................................228 ! List of Tables Table 2.1. Loadings for variables included in principle component analysis. PC1 and PC2 explain 68.32% of the total variance............................................................61 Table 2.2. Reponses (mean ± SE) of grey warbler, fantail and silvereye nest owners to chaffinch (control) and shining cuckoo models at nests......................................62 Table 2.3. PCA score summary (mean ± SE) and values of statistical significance for behavioural differences between model types for grey warblers, fantails and silvereyes. Significant differences are indicated in bold. ....................................63 Table 2.4. Differences among host species in nest defence behaviour in response to a shining cuckoo model or a chaffinch model at the nest. Significant differences are indicated in bold.............................................................................................64 Table 3.1. Red-winged blackbird and gray catbird behaviour at the nest during “pre- cowbird egg” (PRE), “cowbird egg” (CBE), and “post-cowbird egg” (POST) observation periods. Values are reported as means ± SE. P-values < 0.05 were considered significant and are in bold. ................................................................92 Table 3.2. Post-hoc Wilcoxon matched-pairs tests showing the significance of differences between trials where significant differences in gray catbird incubation behaviour at the nest were found among the PRE, CBE and POST trials. P-values < 0.05 were considered significant and are in bold.....................93 Table 3.3. Red-winged blackbird behaviour at the nest according to type of egg manipulation during “pre-cowbird egg” (PRE), “cowbird egg” (CBE) and “post- cowbird egg” (POST) observation periods. Cowbird eggs were either added to blackbird nests, which increased the clutch size (Add) or switched with a host egg, which caused the clutch size to remain the same (Switch). Values are reported as means ± SE........................................................................................94 Table 4.1. Interspecies comparison of gas chromatography (GC-FID) traces of the preen wax of nine species sampled at Kowhai and Waimangara Bushes, South Island, New Zealand. GC-FID traces have not yet been aligned among species but will be aligned once mass spectrometry