Assessing the Impact of Human Disturbance on Penguins
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ASSESSING THE IMPACT OF HUMAN DISTURBANCE ON PENGUINS URSULA ELLENBERG Photo: Hermann Ellenberg, Sandfly Bay, 2007 A thesis submitted for the degree of Doctor of Philosophy at the University of Otago, Dunedin, New Zealand January 2009 Meinen Eltern Gunde und Hermann Ellenberg Assessing the impact of human disturbance on penguins Abstract Nature conservation places great hopes on ecotourism, yet despite increasing pressure on the Earth’s last wild areas, there is insufficient knowledge about human disturbance impact, and hence, visitor management must often remain inadequate. Penguins are an important tourism attraction. Lack of overt behavioural responses to human proximity at breeding sites has led to the common public misconception that penguins would be little affected by human visitation. However, even a single one-off disturbance or a series of low-key disturbances can have far-reaching impacts that are often not immediately discernable to the people involved. I studied behavioural and physiological stress responses of three penguin species that according to anecdotal evidence respond very differently to human proximity. While behaviour could give an indication of the relative severity of a disturbance stimulus, measuring physiological responses was a more objective and powerful tool to quantify stress in all three species studied. As accumulating effects of human disturbance may affect breeding populations, I also monitored a range of reproductive parameters at sites that provided similar breeding habitat and overlapping foraging grounds yet were exposed to different levels of human visitation. Human disturbance can be interpreted as perceived predation risk. Similar to antipredator responses, reaction to disturbance stimuli can affect individual fitness via energetic and lost opportunity costs of risk avoidance. I demonstrated that human proximity is energetically costly for penguins and more so than most stimuli that naturally occur at their breeding sites. Yet the severity of the same disturbance stimulus is perceived differently between species, populations and even individuals. Snares penguins, Eudyptes robustus, appear to be little affected by human activity outside their immediate breeding colony. In comparison, Humboldt penguins, Spheniscus humboldti, proved to be very sensitive to human disturbance. They responded to a person vii Abstract visible at more than 150 m distance from their nest and needed up to half an hour to recover from even careful human approach. Nest densities were lower and breeding success was significantly reduced at more frequently visited sites. In Yellow-eyed penguins, Megadyptes antipodes, I found elevated hormonal and heart rate responses at sites exposed to unregulated visitor access associated with reduced breeding success and lower fledgling weights, which result in reduced first-year survival and recruitment probabilities. Contrary to my expectations, birds that continued to breed despite frequent disturbance had not habituated to human proximity but appeared to be sensitised and responded more strongly to human disturbance. Hence, tourist-exposed penguins were not only disturbed more often, each disturbance event was more costly for the affected birds, eventually leading to poorer breeding performance. Both initial stress response and habituation potential of Yellow-eyed penguins depended on sex, character and previous experiences with humans, and the observed individual differences in behavioural and physiological stress responses related to fitness parameters. Birds characterised as “frozen” were apparently best able to cope in human-disturbed habitats. Thus, human disturbance may drive contemporary evolutionary change in the composition of breeding populations with as yet unrealised consequences for conservation. I give a detailed account of the current understanding of human disturbance related effects on penguins and list future research needs. Complex secondary effects of human disturbance and multiple stressor interactions are still little understood and require long- term population studies that enable us to disentangle human disturbance effects from environmental variability. Similarly, we still know very little about what factors drive habituation or sensitisation in wild populations, thus habituation to even apparently minor human disturbance cannot be assumed. Subtle costs of what we think is minor disturbance can accumulate and may ultimately lead to lower nest numbers, poorer breeding success or reduced fledgling weights in viii Assessing the impact of human disturbance on penguins more frequently visited penguin breeding areas. As the growth of nature-based tourism is expected to continue, it is important not only for ecological but also for economic sustainability to minimise any associated negative human impacts. It is clear that generic guidelines for managing human visitation of penguin colonies cannot be applied. Only rigorous research can provide the basis for appropriate site- and species-specific visitor management decisions. I hope this study can provide a framework for better understanding human disturbance effects on penguins and maybe even on wildlife in general. ix Acknowledgements Acknowledgements A big thanks to Philip Seddon, the best supervisor in the world: You gave me the support and trust I needed to complete this work. Your advice was valuable, feedback immediate and helpful… whether I got knocked down by Murphy’s Law in the field, encountered problems while writing-up or during publishing when having to respond an all-too emotional reviewer. You always were down-to-earth, objective and clear, your suggestions informed and well-thought-off. Thank you for your spontaneous hug when I came to you to admit I became pregnant during the first year of my studies. Phil, I could not have wished for a better supervisor and I am really glad we’ve met and connected. Lloyd Davis, my 10% but much more supervisor: thanks for your humour, good spirits, and the occasional pat on the back. Thank you for the opportunity to join your Snares research team, for shared adventures during a reki trip to Breaksea Island, and for battling our way through silly politics. I once said you “taught me how to talk” and you did – I learned a lot just by observing how brilliantly you communicate. Meinen Eltern Gunde und Hermann Ellenberg gebührt der größte Dank! Ihr habt immer an mich geglaubt und mich in allen Lebenslagen mental wie tatkräftig unterstützt. Ohne euch wäre ich nicht ich und wo ich bin. Eine dicke Umarmung und ganz herzlichen Dank für alles, Ihr könnt euch gar nicht vorstellen wie viel ihr mir bedeutet… Ohne die liebe Omi mit immer neue Ideen, ihre Enkel zu begeistern und zu unterhalten, und den geduldigen Opa, der sich die Zeit nahm für viele Ausflüge ins Museum, an den Strand oder in den Botanischen Garten wäre diese Arbeit auch rein praktisch kaum zu schaffen gewesen. Danke auch für interessiertes und kritisches Korrekturlesen, Schulterklopfen („…also von mir bekämst du eine eins!“), Lübecker Marzipan, Sylter Fischpfanne und Königskuchen! All my love to Thomas Mattern, my partner, best friend, and father of our two delightful kids. We are a great team in the field and surprisingly resistant to daily grind. Much of this work happened side by side with you. Thank you for your support, whether hands on during fieldwork, philosophically when discussing great ideas or concerns past midnight, x Assessing the impact of human disturbance on penguins or during last minute formatting or proof reading. You grounded me in reality when my optimism made me soar high up in the clouds and you pulled me up when I was weighed down by worries. I am glad to have you, Thomas, and I enjoy surfing together with you on top of everyday challenges. Hannah Lea and Finn-Erik, you are my life. I cannot imagine how people manage to write a PhD without kids. With you I had to be organised and then could discover the pleasures of slowness. You helped me to ignore the pressure of looming deadlines and enjoy the really important things in life: The sickle of the moon over the busy parking lot of a busy supermarket after a busy day, the crawly bugs under a rock that are busy too, splashing through the waves during a stroll along a rainy winter’s beach, smelling a meadow full of flowers, singing a song and dancing along, discovering gnome hide-outs in the cool of the summer forest, flying a kite high up into the clouds, picking strawberries warmed by the sun or huddling together with a nice story book in front of our fireplace. You understood when I was tired after a short night and made sure I never slept in. Finn, you keen observer with great imagination and persuasion… I liked your “Let’s stay a bit longer at the beach and give the fairies some more time to finish your PhD”, your rainbow dinosaur when I felt low, your pirate treasure hunts, castles and knights, stories of Maui, your tireless energy, your challenging “why”-questions, air-guitar play and improvised songs. Hannah: “maam – lala!” your love for music and dance is good for the soul, your smile and happiness very infectious. Thank you for patient proof reading, uplifting cuddles, and clearly communicating priorities. Thank you both for your trust – I hope I will never let you down. A big cuddle and thank you to our wee new flatmate Ylva Nele! You waited patiently in my tummy while I was doing my revisions and until your siblings could not wait any longer and called excitedly: “Come out you sleepy head - we want to meet you!” My brother Lars-Gunnar “…do you have a problem for me to solve? …I am a mind worker – I need a problem, now!” is a great resource when you are stuck in ideas but lack the ability to put these into code. Thank you for your patience and immediate response solving xi Acknowledgements many silly computer problems. You programmed the heart rate analysis software just for fun back in the days when I worked on Arctic terns and adapted scripts for penguins.