The maintenance of both the psychological and physiological health of captive animals is a key priority of modern zoos. Recognising that characteristics of the captive environment have the potential to decrease animal welfare, methods for quantifying and assessing welfare have been developed as part of the process for improving animal welfare. Traditionally, observations of animal behaviour and quantifying time budgets in relation to those of the animals’ wild counterparts have been utilised to assess animal welfare. Hormonal assays have also been implemented to quantify the physiological stress response of animals in captivity and identify the extent of stress being experienced. Each of these methods focuses on a different indicator of animal welfare, is quantified in different ways and provides a different perspective on the welfare of the animals. Given the limited time and financial budgets available to zoos and animal carers, identifying the most appropriate method of welfare assessment would be advantageous in helping to secure the best possible health of captive animals and to maximise their value in captivity. This thesis implemented both behavioural observations and hormonal assays to identify the strengths and weaknesses of each methodology, and make recommendations for future research. The study involved a group of four Bornean orangutans (Pongo pygmaeus) housed at Twycross Zoo. Behavioural observations involved continuous group sampling and the development of an ethogram to record a comprehensive account of orangutan activity over the course of a 12 week enrichment programme. Simultaneous to these observations, faecal samples were collected from each orangutan and processed via Enzyme Immunoassay (EIA) to quantify levels of faecal glucocorticoid metabolites (fGCM) in each sample. While recognising the recent developments in ecological analytical methods, the capacity for extending network analysis beyond the application to social networks, and its use as a welfare assessment tool were explored. Behavioural and space-use networks were developed using data from a second study of the orangutans housed at Twycross Zoo. The flexibility of network analysis in visually representing different data types allowed for the intuitive representation of complex behavioural data. Further research investigated the use of network metrics in providing deeper insights into animal behaviour and space use patterns. In addition, bipartite networks were assessed for their potential to detect and show patterns in the relationships between two sets of behavioural data. Each of the methods used had a number of strengths and weaknesses, but importantly each contributed a different perspective in the assessment of behaviour patterns and welfare, suggesting that an integrated approach to behaviour studies utilising several methods would be ideal. Cost and logistic constraints make this unlikely in most cases. However, the thesis ends with a look to the future and the recognition that the current rapid development of technology for use in animal behaviour studies, coupled with equally rapid development of analytical techniques, may help to dramatically increase the amount of information gained from the average animal behaviour study in the future. Such improvements have never been more urgent, with the requirement for understanding animal behaviour in light of current extinction rates within the context of habitat destruction and climate change. It is hoped that this thesis will make a contribution to improving future animal behaviour and welfare studies by providing an assessment of both traditional methods of study as well as demonstrating the use and potential of new ways of applying network analysis within such studies.

Migration timing is a conserved life-history trait. To address the hypothesis that reproductive hormones are principal determinants of migration timing, I physiologically biopsied over 1000 sockeye salmon and monitored their subsequent behaviour with acoustic and radio telemetry as they migrated from the Pacific Ocean toward and into the Fraser River, and then onward to distant spawning areas. Links between physiology, behaviour, and survival were examined. Circulating testosterone was found to be positively correlated with the rates of river entry in Late-run females but not in males, despite having concentrations that were equal if not higher than those of females. The notion of protandrous migration, in which males synchronize their activities to the reproductive and migratory schedules of females, was postulated as the basis for this difference. Once in river however, successful males and females were those that (1) took longest to enter the river, and (2) had high somatic energy, low testosterone, and low gill Na+,K+-ATPase activities. An experimental test of the effect of reproductive hormones on the regulation of migration timing proved inconclusive. Relative to controls, GnRH and (or) testosterone treatment did not influence rates of ocean travel by males. Unfortunately, no females were examined. Nevertheless, significant, positive correlations between initial testosterone and travel times were found irrespective of hormonal treatment, which was unexpected but consistent with the previous studies. In an experimental simulation of an ‘early’ migration, normally timed Late-run sockeye exposed to typical 10 ºC river temperatures and then released to complete migration were 68% successful. In contrast, salmon held at 18 °C and released were half as successful. The expression of a kidney parasite was near maximal in the 18 °C fish and undetectable in the 10 °C fish. Only gill Na+,K+-ATPase activity differed between groups, with a drop in the 18 °C fish. Though no clear stress, reproductive, or energetic differences were observed between groups, the ultimate effect of high temperature treatment was high disease expression, slowed migration speeds, and high migration mortality. Changes in reproductive schedules, due to changes in latitudinal ocean distributions, are discussed as potential causes of early migration by Late-run sockeye.

Migration timing is a conserved life-history trait. To address the hypothesis that reproductive hormones are principal determinants of migration timing, I physiologically biopsied over 1000 sockeye salmon and monitored their subsequent behaviour with acoustic and radio telemetry as they migrated from the Pacific Ocean toward and into the Fraser River, and then onward to distant spawning areas. Links between physiology, behaviour, and survival were examined. Circulating testosterone was found to be positively correlated with the rates of river entry in Late-run females but not in males, despite having concentrations that were equal if not higher than those of females. The notion of protandrous migration, in which males synchronize their activities to the reproductive and migratory schedules of females, was postulated as the basis for this difference. Once in river however, successful males and females were those that (1) took longest to enter the river, and (2) had high somatic energy, low testosterone, and low gill Na+,K+-ATPase activities. An experimental test of the effect of reproductive hormones on the regulation of migration timing proved inconclusive. Relative to controls, GnRH and (or) testosterone treatment did not influence rates of ocean travel by males. Unfortunately, no females were examined. Nevertheless, significant, positive correlations between initial testosterone and travel times were found irrespective of hormonal treatment, which was unexpected but consistent with the previous studies. In an experimental simulation of an ‘early’ migration, normally timed Late-run sockeye exposed to typical 10 ºC river temperatures and then released to complete migration were 68% successful. In contrast, salmon held at 18 °C and released were half as successful. The expression of a kidney parasite was near maximal in the 18 °C fish and undetectable in the 10 °C fish. Only gill Na+,K+-ATPase activity differed between groups, with a drop in the 18 °C fish. Though no clear stress, reproductive, or energetic differences were observed between groups, the ultimate effect of high temperature treatment was high disease expression, slowed migration speeds, and high migration mortality. Changes in reproductive schedules, due to changes in latitudinal ocean distributions, are discussed as potential causes of early migration by Late-run sockeye.

Migration timing is a conserved life-history trait. To address the hypothesis that reproductive hormones are principal determinants of migration timing, I physiologically biopsied over 1000 sockeye salmon and monitored their subsequent behaviour with acoustic and radio telemetry as they migrated from the Pacific Ocean toward and into the Fraser River, and then onward to distant spawning areas. Links between physiology, behaviour, and survival were examined. Circulating testosterone was found to be positively correlated with the rates of river entry in Late-run females but not in males, despite having concentrations that were equal if not higher than those of females. The notion of protandrous migration, in which males synchronize their activities to the reproductive and migratory schedules of females, was postulated as the basis for this difference. Once in river however, successful males and females were those that (1) took longest to enter the river, and (2) had high somatic energy, low testosterone, and low gill Na+,K+-ATPase activities. An experimental test of the effect of reproductive hormones on the regulation of migration timing proved inconclusive. Relative to controls, GnRH and (or) testosterone treatment did not influence rates of ocean travel by males. Unfortunately, no females were examined. Nevertheless, significant, positive correlations between initial testosterone and travel times were found irrespective of hormonal treatment, which was unexpected but consistent with the previous studies. In an experimental simulation of an ‘early’ migration, normally timed Late-run sockeye exposed to typical 10 ºC river temperatures and then released to complete migration were 68% successful. In contrast, salmon held at 18 °C and released were half as successful. The expression of a kidney parasite was near maximal in the 18 °C fish and undetectable in the 10 °C fish. Only gill Na+,K+-ATPase activity differed between groups, with a drop in the 18 °C fish. Though no clear stress, reproductive, or energetic differences were observed between groups, the ultimate effect of high temperature treatment was high disease expression, slowed migration speeds, and high migration mortality. Changes in reproductive schedules, due to changes in latitudinal ocean distributions, are discussed as potential causes of early migration by Late-run sockeye. / Forestry, Faculty of / Graduate