Measuring hormonal changes is vital for understanding how the social and physical environment influences behaviour, reproduction and survival. Various methods of hormone measurement exist, potentially explaining variation in results across studies; methods should be cross validated to ensure they correlate. I directly compare faecal and plasma hormone measurements (Chapter 2), and use the most suitable endocrine measure to test the Darling hypothesis (Chapter 3) - that breeding is hastened and synchronized in larger colonies due to increased social stimulation (mediated by the endocrine system). Blood and faecal samples were simultaneously collected from individual Adelie penguins (Pygoscelis adeliae) for comparison, and assayed for testosterone and corticosterone (or their metabolites). Sex differences and variability within each measure, and correlation of values across measures were compared. For both hormones, plasma samples showed greater variation than faecal samples. Males had higher corticosterone levels than females, but the difference was only significant in faecal samples. Plasma testosterone, but not faecal testosterone, was significantly higher in males than females. Correlation between sample types was poor overall, and weaker in females than in males; perhaps because measures from plasma represent hormones that are both free and bound to globulins, whereas measures from faeces represent only the free portion. Faecal samples also represent a cumulative measure of hormones over time, as opposed to a plasma 'snapshot' concentration. Faecal sampling appears more suitable for assessing baseline hormone levels. In the second study I examined, over two seasons, whether the timing of breeding varied with colony size; larger colonies present occupants with higher levels of social stimulation and are predicted to show earlier, more synchronous breeding. Baseline faecal hormone levels throughout the breeding season, and survival, were measured to investigate possible proximate and ultimate mechanisms for the results. The influence of environmental variability was examined, by relating the timing of breeding, survival, and endocrine changes to sea ice conditions. Colony size did not influence the timing or synchrony of breeding, survival, or hormone levels within years; perhaps because colonies in an Adelie rookery are not independent from the 'social environment' of adjacent colonies. Across years, synchrony in the smaller rookery was higher than in the larger rookery. The scale of these comparisons may exceed the applicability of the Darling hypothesis. Therefore, no support was found for the Darling hypothesis, at the colony or rookery level, in this species. Higher corticosterone metabolite and lower sex hormone levels in the first season correlated to later breeding and lower survival compared to the second season. This is likely due to the persistence of extensive sea ice conditions late into the first season. Researchers should take care in selecting the most appropriate method of hormone measurement for their question. Future studies testing the Darling hypothesis must carefully select their definition of a colony (i.e. a truly isolated social unit) and the scale at which the hypothesis is tested. Combining endocrine measurements with behavioural, survival, and environmental information allows for a more comprehensive interpretation of animal ecology.

The ability to understand the complex mechanisms by which species communicate has been the focus of study for many years. The Adélie penguin (Pygoscelis adeliae) breeds in vocally challenging Antarctic colonies and has evolved a complex system of intra- and intersexual visual and vocal behaviours. Unique to the Adélie penguin, is the presence of two distinct display calls, the Loud Mutual Display (LMD) call and male Ecstatic Display Call (EDC). The EDC is used for both mate choice and territory defence; but apart from general structure and function this call remains basically unstudied. Therefore, the aim of this study is to examine the EDC with regard to its function (during mate choice) and variability over time, with location and with changing conditions. It is predicted that the EDC will be an individual call that is distinct from the LMD call. Additionally, the effects of an inclement breeding season on factors that affect breeding success will be examined using a logistic regression approach. The study took place over two breeding seasons(2000/1 & 2002/3) on Ross Island, Antarctica, with the majority of research being conducted at Cape Bird. Results of the study indicate that the EDC is an individual call that cannot reliably be described as a male display as females also give an EDC that is acoustically comparable. The similarities between the EDC and the LMD call seem to relate only to syllabic organisation (structure), as almost every call parameter was different between call types. Furthermore, several call parameters differed between Ross Island colonies. These parameters varied with both geographical location and colony size. In conjunction with variation in call parameters, the health and breeding success of the colonies was different. The stability of call parameters (from Cape Bird) indicated that parameters that separate colonies not only alter between breeding years, but also vary during a single breeding season. The function of the EDC as an honest signal during mate choice was supported by spectral analyses. These analyses found heavier males had significantly lower Frequency Modulation (FM) in the short repeated syllables of their calls than lighter males. Furthermore, a male’s breeding success could be predicted during the early breeding season by the FM. Logistic regression models showed that early male arrival, weight and good nest quality were most influential in predicting fledging success. The logistic regression method also showed that the parameters measured varied in their predictive ability as the season progressed. This study has not only enhanced the wealth of knowledge regarding Adélie breeding behaviour and developed our understanding of the vocal repertoire of this species, but also has implications for honest signalling theory, and the evolution and stability of vocal variation at different spatial and temporal scales.

The ability to understand the complex mechanisms by which species communicate has been the focus of study for many years. The Ad��lie penguin (Pygoscelis adeliae) breeds in vocally challenging Antarctic colonies and has evolved a complex system of intra- and intersexual visual and vocal behaviours. Unique to the Ad��lie penguin, is the presence of two distinct display calls, the Loud Mutual Display (LMD) call and male Ecstatic Display Call (EDC). The EDC is used for both mate choice and territory defence; but apart from general structure and function this call remains basically unstudied. Therefore, the aim of this study is to examine the EDC with regard to its function (during mate choice) and variability over time, with location and with changing conditions. It is predicted that the EDC will be an individual call that is distinct from the LMD call. Additionally, the effects of an inclement breeding season on factors that affect breeding success will be examined using a logistic regression approach. The study took place over two breeding seasons(2000/1 & 2002/3) on Ross Island, Antarctica, with the majority of research being conducted at Cape Bird. Results of the study indicate that the EDC is an individual call that cannot reliably be described as a male display as females also give an EDC that is acoustically comparable. The similarities between the EDC and the LMD call seem to relate only to syllabic organisation (structure), as almost every call parameter was different between call types. Furthermore, several call parameters differed between Ross Island colonies. These parameters varied with both geographical location and colony size. In conjunction with variation in call parameters, the health and breeding success of the colonies was different. The stability of call parameters (from Cape Bird) indicated that parameters that separate colonies not only alter between breeding years, but also vary during a single breeding season. The function of the EDC as an honest signal during mate choice was supported by spectral analyses. These analyses found heavier males had significantly lower Frequency Modulation (FM) in the short repeated syllables of their calls than lighter males. Furthermore, a male���s breeding success could be predicted during the early breeding season by the FM. Logistic regression models showed that early male arrival, weight and good nest quality were most influential in predicting fledging success. The logistic regression method also showed that the parameters measured varied in their predictive ability as the season progressed. This study has not only enhanced the wealth of knowledge regarding Ad��lie breeding behaviour and developed our understanding of the vocal repertoire of this species, but also has implications for honest signalling theory, and the evolution and stability of vocal variation at different spatial and temporal scales.

The ability to understand the complex mechanisms by which species communicate has been the focus of study for many years. The Adélie penguin (Pygoscelis adeliae) breeds in vocally challenging Antarctic colonies and has evolved a complex system of intra- and intersexual visual and vocal behaviours. Unique to the Adélie penguin, is the presence of two distinct display calls, the Loud Mutual Display (LMD) call and male Ecstatic Display Call (EDC). The EDC is used for both mate choice and territory defence; but apart from general structure and function this call remains basically unstudied. Therefore, the aim of this study is to examine the EDC with regard to its function (during mate choice) and variability over time, with location and with changing conditions. It is predicted that the EDC will be an individual call that is distinct from the LMD call. Additionally, the effects of an inclement breeding season on factors that affect breeding success will be examined using a logistic regression approach. The study took place over two breeding seasons(2000/1 & 2002/3) on Ross Island, Antarctica, with the majority of research being conducted at Cape Bird. Results of the study indicate that the EDC is an individual call that cannot reliably be described as a male display as females also give an EDC that is acoustically comparable. The similarities between the EDC and the LMD call seem to relate only to syllabic organisation (structure), as almost every call parameter was different between call types. Furthermore, several call parameters differed between Ross Island colonies. These parameters varied with both geographical location and colony size. In conjunction with variation in call parameters, the health and breeding success of the colonies was different. The stability of call parameters (from Cape Bird) indicated that parameters that separate colonies not only alter between breeding years, but also vary during a single breeding season. The function of the EDC as an honest signal during mate choice was supported by spectral analyses. These analyses found heavier males had significantly lower Frequency Modulation (FM) in the short repeated syllables of their calls than lighter males. Furthermore, a male’s breeding success could be predicted during the early breeding season by the FM. Logistic regression models showed that early male arrival, weight and good nest quality were most influential in predicting fledging success. The logistic regression method also showed that the parameters measured varied in their predictive ability as the season progressed. This study has not only enhanced the wealth of knowledge regarding Adélie breeding behaviour and developed our understanding of the vocal repertoire of this species, but also has implications for honest signalling theory, and the evolution and stability of vocal variation at different spatial and temporal scales.

The ability to understand the complex mechanisms by which species communicate has been the focus of study for many years. The Adélie penguin (Pygoscelis adeliae) breeds in vocally challenging Antarctic colonies and has evolved a complex system of intra- and intersexual visual and vocal behaviours. Unique to the Adélie penguin, is the presence of two distinct display calls, the Loud Mutual Display (LMD) call and male Ecstatic Display Call (EDC). The EDC is used for both mate choice and territory defence; but apart from general structure and function this call remains basically unstudied. Therefore, the aim of this study is to examine the EDC with regard to its function (during mate choice) and variability over time, with location and with changing conditions. It is predicted that the EDC will be an individual call that is distinct from the LMD call. Additionally, the effects of an inclement breeding season on factors that affect breeding success will be examined using a logistic regression approach. The study took place over two breeding seasons(2000/1 & 2002/3) on Ross Island, Antarctica, with the majority of research being conducted at Cape Bird. Results of the study indicate that the EDC is an individual call that cannot reliably be described as a male display as females also give an EDC that is acoustically comparable. The similarities between the EDC and the LMD call seem to relate only to syllabic organisation (structure), as almost every call parameter was different between call types. Furthermore, several call parameters differed between Ross Island colonies. These parameters varied with both geographical location and colony size. In conjunction with variation in call parameters, the health and breeding success of the colonies was different. The stability of call parameters (from Cape Bird) indicated that parameters that separate colonies not only alter between breeding years, but also vary during a single breeding season. The function of the EDC as an honest signal during mate choice was supported by spectral analyses. These analyses found heavier males had significantly lower Frequency Modulation (FM) in the short repeated syllables of their calls than lighter males. Furthermore, a male’s breeding success could be predicted during the early breeding season by the FM. Logistic regression models showed that early male arrival, weight and good nest quality were most influential in predicting fledging success. The logistic regression method also showed that the parameters measured varied in their predictive ability as the season progressed. This study has not only enhanced the wealth of knowledge regarding Adélie breeding behaviour and developed our understanding of the vocal repertoire of this species, but also has implications for honest signalling theory, and the evolution and stability of vocal variation at different spatial and temporal scales.

The ability to understand the complex mechanisms by which species communicate has been the focus of study for many years. The Adélie penguin (Pygoscelis adeliae) breeds in vocally challenging Antarctic colonies and has evolved a complex system of intra- and intersexual visual and vocal behaviours. Unique to the Adélie penguin, is the presence of two distinct display calls, the Loud Mutual Display (LMD) call and male Ecstatic Display Call (EDC). The EDC is used for both mate choice and territory defence; but apart from general structure and function this call remains basically unstudied. Therefore, the aim of this study is to examine the EDC with regard to its function (during mate choice) and variability over time, with location and with changing conditions. It is predicted that the EDC will be an individual call that is distinct from the LMD call. Additionally, the effects of an inclement breeding season on factors that affect breeding success will be examined using a logistic regression approach. The study took place over two breeding seasons(2000/1 & 2002/3) on Ross Island, Antarctica, with the majority of research being conducted at Cape Bird. Results of the study indicate that the EDC is an individual call that cannot reliably be described as a male display as females also give an EDC that is acoustically comparable. The similarities between the EDC and the LMD call seem to relate only to syllabic organisation (structure), as almost every call parameter was different between call types. Furthermore, several call parameters differed between Ross Island colonies. These parameters varied with both geographical location and colony size. In conjunction with variation in call parameters, the health and breeding success of the colonies was different. The stability of call parameters (from Cape Bird) indicated that parameters that separate colonies not only alter between breeding years, but also vary during a single breeding season. The function of the EDC as an honest signal during mate choice was supported by spectral analyses. These analyses found heavier males had significantly lower Frequency Modulation (FM) in the short repeated syllables of their calls than lighter males. Furthermore, a male’s breeding success could be predicted during the early breeding season by the FM. Logistic regression models showed that early male arrival, weight and good nest quality were most influential in predicting fledging success. The logistic regression method also showed that the parameters measured varied in their predictive ability as the season progressed. This study has not only enhanced the wealth of knowledge regarding Adélie breeding behaviour and developed our understanding of the vocal repertoire of this species, but also has implications for honest signalling theory, and the evolution and stability of vocal variation at different spatial and temporal scales.