Conservation Ecology and Breeding Biology of the White-Browed Treecreeper Climacteris Affinis
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Conservation Ecology and Breeding Biology of the White-browed Treecreeper Climacteris affinis James Radford B.Sc. (Hons) University of Melbourne A thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy of Deakin University June 2002 Acknowledgments First and foremost, I would like to thank my supervisor, Andrew Bennett, for his tireless encouragement, enthusiasm, direction and wisdom throughout my candidature. Your door was always open and your advice re-assuring. Thanks AB. This research was possible because I was the recipient of an Australian Postgraduate Award. I would also like to acknowledge the financial assistance provided by the Department of Natural Resources and Environment (North-west region), the School of Ecology and Environment at Deakin University, Birds Australia through the auspices of the Stuart Leslie Bird Research Award, the M. A. Ingram Trust and the Society for Conservation Biology (travel award). Drs. Ralph MacNally (Monash University) and John White (Deakin University) generously offered advice on statistical analysis and other matters of research design. Chris Lewis and Rowena Scott (Deakin University) provided invaluable assistance with technical, logistical and day-to-day business and their help was greatly appreciated. Jackie MacReadie, Andrew Giles and Grant Michie ensured my administrative affairs were in order. My thanks to Charles and Joan Sandbrink, Kahlid Al-Dabbagh, and the banding team at Rushworth for their assistance with my bird-banding training. I received great support and assistance from Geoffrey Allen and Victor Hurley (Flora and Fauna, NRE, Mildura). Special thanks to all the Parks Victoria staff at Werrimull and Wyperfeld, especially Lorraine Ludewigs, Peter Teasdale, Jack Kelly, Travis Harman and Brian Goldsmith. You can have your ladder back now! Many people helped in the field, either practically with data collection, or just as importantly, with their company and good cheer. In order of appearance, my thanks to John Bardsley, Tim Bardsley, Wendy Radford, Geoffrey Allen, Alec Hawtin, John Morrish, Mary Conway, Barry McLean, Victor Hurley, Sue-Ellen and Georgia Radford, Megan Benne, James Kilby, Ken Hudson, Helen Bardsley, Kate Hudson, Anne Balfe, Peter Balfe, Grant Palmer and Ben Morris. Special thanks to Tim and John for their repeat-appearances, and to Alec for his sound recordings of the White- browed Treecreeper. i My thanks to the following landholders for permission to search for treecreepers on their property: Greg and Toni Arney, Rod Rowe, Lindsay and Anne Morrish, Neil and Jenny Boord, Jack Sutherland, Russell Abraham, Ray and Janine Frances, Don and Caroline Rankin. I am especially grateful to Janine and Ray, and Caroline and Don for their hospitality, meals, welcoming smiles and the occasional cold beer when it was most needed. Thanks. I was not studying the treecreepers alone! My thanks to the other members of the “Treecreeper mafia”, Gary Luck, Veronica Doerr and Erik Doerr, who willing gave of their expertise in all matters Climacteridae. Gary Luck and Richard Retallick provided valuable comments on draft chapters. The post-grad students of the School of Ecology and Environment provided support and good humour, and I would especially like to thank the other members of the Landscape Ecology Research Group: Rodney van der Ree, Jenny Wilson (you did get there first Jen, but only just!), Chris Tzaros, Grant Palmer, Mark Antos, Lindy Lumsden, Greg Holland and Lindy Banks. I am deeply grateful for the moral support, love and interest (not to mention the scope!) provided by my family during the arduous course of this thesis. To Mum, John, Tim and Lou, my heartfelt thanks. This research has presented many challenges and hardships. None more so than the weeks on end I was required to spend away from my wife and daughter, whose birth coincided with the commencement of my first field season. Having managed to navigate through those difficult times, we struck the seemingly endless hours of analysis and writing. Suze, I am forever grateful for your commitment to our common goals, courage, understanding (although it was tested at times) and love throughout this process. Georgia, my sweet, Dad has finished working now! ii Abstract The White-browed Treecreeper Climacteris affinis is one of many woodland- dependent birds that are at risk from the encroachment of human-dominated land- uses into natural landscapes. The White-browed Treecreeper inhabits semi-arid woodlands in north-west Victoria, Australia, a vegetation community that has undergone extreme modification in the last century due to the expansion of agriculture in the region. Extant woodlands represent only 10% of the original woodland cover in the region, and are highly fragmented and disturbed in many districts. Thus, the survival of the White-browed Treecreeper may depend on active management. However, current knowledge of the ecology and biology of this species is virtually non-existent, and inadequate for informed and effective conservation actions. The aim of this thesis is to redress this situation and provide the ecological basis for sound conservation management of the species. The thesis consists of two parts: an investigation of habitat use at three spatial scales and a study of the social organization, nesting requirements, breeding behaviour and reproductive success of a population of White-browed Treecreepers. Fifty-six patches of remnant woodland in north-west Victoria were surveyed to determine the factors affecting the occurrence of the White-browed Treecreeper at the regional scale. It was detected in 16 patches, and was largely confined to two core districts - Yarrara and Wyperfeld (Pine Plains). The floristic composition of the dominant tree species was an important determinant of patch occupancy, with the results providing quantitative support for the previously suspected affinity for Belah Casuarina pauper and Slender Cypress-pine Callitris gracilis – Buloke Allocasuarina luehmannii woodlands. However, the absence of the White-browed Treecreeper from several districts was due to factors other than a lack of appropriate habitat. Demographic isolation – the distance from the focal patch to the nearest population of the White-browed Treecreeper – was the most important variable in explaining variation in patch occupancy. Patches isolated from other treecreeper populations by more than 8.3 km in landscapes of non-preferred native vegetation, and 3 km in agricultural landscapes, were unlikely to support the White-browed Treecreeper. The impact of habitat loss and fragmentation on the capacity of individuals to move through the landscape (i.e. functional connectivity) is considered in relation to disruption to dispersal and migration, and the potential collapse of local metapopulations. iii Habitat use was then examined in a network of patches and linear strips of Belah woodland embedded in a predominantly cultivated landscape. A minimum area of 18.5 ha of Belah woodland was identified as the most important criterion for patch occupancy at the local scale. This landscape appeared to be permeable to movement by the White-browed Treecreeper, facilitated by the extensive network of linear habitat, and clusters of small to medium fragments. The third scale of habitat use investigated the frequency of use of 1-ha plots within tracts of occupied woodland. It is important to discriminate between habitat traits that operate at the population level, and those that act as proximate cues for habitat selection by individuals. Woodlands that have high tree density, extensive cover of low-stature shrubs, abundant lichen, a complex vertical structure, and relatively low cover of grass and herbs are likely to support larger populations of the White-browed Treecreeper. However, individuals appeared to be using tree dominance (positive) and tall shrub cover (negative) as proximate environmental stimuli for habitat selectivity. A relatively high cover of ground lichen, which probably reflects a ground layer with low disturbance and high structural complexity, was also a reliable indicator of habitat use. Predictive models were developed which could be used to plan vegetation management to enhance habitat for the White-browed Treecreeper. The results of the regional, landscape and patch-scale investigations emphasise that factors operating at multiple spatial scales influence the suitability of remnant vegetation as habitat for the White-browed Treecreeper. The White-browed Treecreeper is typical of many small Australian passerines in that it has high annual survival, small clutches, a long breeding season, multiple broods and relatively low reproductive rates. Reproductive effort is adjusted through the number of clutches laid rather than clutch size. They occupy relatively large, all- purpose territories throughout the year. However, unlike many group territorial birds, territory size was not related to the number of occupants. The White-browed Treecreeper nests in tree hollows. They select hollows with a southerly orientation where possible, and prefer hollows that were higher from the ground. At Yarrara, there was considerable spatial variation in hollow abundance that, in concert with territorial constraints, restricted the actual availability of hollows to less than the absolute abundance of hollows. Thus, the availability of suitable hollows may limit reproductive productivity in some territories, although the iv magnitude of this constraint on overall population growth is predicted