Tall eucalypt forests of Australia: structure, pattern and process TALL EUCALYPT FORESTS OF AUSTRALIA: STRUCTURE, PATTERN AND PROCESS Jessie C. Buettel B.Sc. (Hons) UTas School of Biological Sciences – University of Tasmania TAS 7001 AUSTRALIA Thesis submitted in fulfilment of the requirements for the Degree of Doctor of Philosophy Submitted June, 2017 1 Tall eucalypt forests of Australia: structure, pattern and process 2 Tall eucalypt forests of Australia: structure, pattern and process DECLARATION This thesis contains no material which has been accepted for the award of any other degree or diploma in any tertiary institution, and to the best of my knowledge and belief, contains no material previously published or written by another person, except where due reference is made in the text of the thesis, nor does the thesis contain any material that infringes copyright. The publishers of the papers comprising all chapters, hold the copyright for that content, and access to the material should be sought from the respective journal. The remaining non-published content of the thesis may be made available for loan and limited copying and communication in accordance with the Copyright Act 1968. Date Signed 18 June 2017 Jessie C. Buettel 3 Tall eucalypt forests of Australia: structure, pattern and process STATEMENT OF ETHICAL CONDUCT The research associated with this thesis abides by the international and Australian codes on human and animal experimentation, the guidelines by the Australian Government's Office of the Gene Technology Regulator and the rulings of the Safety, Ethics and Institutional Biosafety Committees of the University. Date Signed 18 June 2017 Jessie C. Buettel 4 Tall eucalypt forests of Australia: structure, pattern and process STATEMENT OF PUBLISHED WORK Candidate was the primary author on all chapters, undertook substantial fieldwork, and did most data analyses. For each respective chapter, the candidate as well as all co-authors contributed to developing the ideas. Barry W. Brook and Nicole Bezemer assisted with the field work – collecting fallen wood and topographic data that contributed to Chapters III, VI, VIII and IX. David M. J. S. Bowman provided access to the primary data used in Chapters III, IV, VI, VII and IX. Barry W. Brook, Stefania Ondei, George P. W. Perry, John Alroy, Andrew Cole and John Dickey contributed to some data analysis. The candidate as well as Barry W. Brook assisted with refining the text. Supplementary Information for all chapters, including data and additional analyses, can be found here: https://ecological-dynamics.org/data/ The following papers have resulted from this Ph.D. thesis research: Buettel, J.C., Bowman, D.M.J.S., Perry, G.L.W., Hovenden, M.J. & Brook, B.W. (2016) A global synthesis of reported spatial patterns in forests. Ecography (in review). Buettel, J.C., Perry, G.W.P., Ondei, S. & Brook B.W. (2017) Drivers of spatial pattern, density and basal area in Australian tall eucalypt forests. Journal of Ecology (in review). 5 Tall eucalypt forests of Australia: structure, pattern and process Buettel, J.C., Alroy, J. & Brook, B.W. (2017) Latitudinal controls on forest species richness are not direct. Nature Ecology and Evolution (in review). Buettel, J.C., Ondei, S. & Brook, B.W. (2017) Look down to see what’s up: A systematic overview of treefall dynamics in forests. Forests 8, 123. doi:10.3390/f8040123. Buettel, J.C., Ondei, S., Dickey, J.M., Cole, A.A. & Brook, B.W. (2017) Reading the forest logbook: environmental drivers of living and dead elements of Australian eucalypt forests. Diversity and Distributions (in review). Buettel, J.C., Ondei, S. & Brook, B.W. (2017) Missing the wood for the trees? New ideas on defining forests and forest degradation. Rethinking Ecology (in press). doi: 10.3897/[email protected] Buettel, J.C., Ondei, S. & Brook, B.W. (2017) A practical method for creating a digital topographic surface for ecological plots using ground-based measurements. Landscape Ecology (in review). Buettel, J.C., Cole, A., Dickey, J.M. & Brook, B.W. (2017) Modelling linear spatial features in ecology. Ecology (in review). Buettel, J. C., Brook, B.W., Cole, A.A., Dickey, J.M. & Flies, E.J. (2017) Interdisciplinary to transdisciplinary: shifting the collaboration paradigm for greater advances in science. Ecology and Evolution. (in review). Buettel, J.C. & Brook, B.W. (2016) Egress! How technophilia can reinforce biophilia to improve ecological restoration. Restoration Ecology 24, 843-847. doi: 10.1111/rec.12387 (article featured on the journal cover). 6 Tall eucalypt forests of Australia: structure, pattern and process Additional publications candidate co-authored during PhD: Brook, B.W. & Buettel, J.C. (2016) Emigration is costly but immigration has benefits in human-altered landscapes. Functional Ecology 30, 1478-1479. doi: 10.1111/1365-2435.12639. Brook, B.W., Ellis, E.C. & Buettel, J.C. (2017) What is the evidence for planetary tipping points? Chapter 8 in: Effective Conservation Science: Data Not Dogma (eds. P. Kareiva, M. Marvier & B. Silliman), Oxford University Press (in press). ISBN: 9780198808985. Johnson, C. N., Balmford, A., Brook, B.W., Buettel, J.C., Galetti, M., Guangchun, L. & Wilmshurst, J.M. (2017) Biodiversity losses and conservation responses in the Anthropocene. Science 356, 270-275 (major review). doi: 10.1126/science.aam9317. Signed: Signed: Barry W. Brook Anthony Koutoulis Supervisor Head of School School of Biological Sciences School of Biological Sciences University of Tasmania University of Tasmania Date: 18 June 2016 Date: 18 June 2016 7 Tall eucalypt forests of Australia: structure, pattern and process ACKNOWLEDGEMENTS There are many people that have been part of my journey these last 3.5 years, and the stress of forgetting anyone is almost equivalent to ‘waiting for the in-review status to change to awaiting editor decision for a manuscript submission!’ So, I will start by offering a sincere thank you to everyone that has had an impact on my PhD at any stage throughout my journey – you may not have been mentioned in these acknowledgments but be rest assured that my bad memory is the reason! I would, first and foremost like to give a wholehearted thank you to my supervisor Barry Brook, for taking me under his wing and providing me with ample opportunities to grow and excel. There was never a challenge he thought I couldn’t rise to; his constant belief in my ability, and support and encouragement that he provided will remain with me for the rest of my career. I am proud, and sincerely blessed to have been under his tutelage. My PhD would not have been possible without the AusPlots data and David Bowman’s belief that I would be able to do it justice. TERN (AusPlots), Sam Wood, Karl Rann and everyone associated with the funding and collection of these data – I am deeply grateful. A massive thank you to Nicole Bezemer for her dedication and help with field work. We created some great memories on those trips. John Dickey and Andrew Cole – thank you for your enthusiasm and willingness to dabble in ecology, and providing me with support and encouragement along the way. I never thought I would have had such stimulating conservations about the analogies between galaxies (stars) and ecology (trees). 8 Tall eucalypt forests of Australia: structure, pattern and process Jon, I am deeply grateful for your friendship, guidance and support throughout this journey, I could not have done it without you. I also wish to thank my close friends and family for their support and patience throughout this journey. My family’s mantra: ‘you are Jessie Buettel and you can do anything,’ is a perfect testament to their unwavering belief in me. Stefania Ondei, you are amazing. That is all. To the D.E.E.P lab – I appreciate your patience and encouragement during my PhD completion. You are a wonderful and fun group of people. Sam Adlard, Ian Cover, Emily Flies, thank you for being there for me and believing in me – particularly through all the tough times. A warm thanks to Scott Carver, Vector, Relly, (the puppas) and Lynx for being my rock(s) through turbulent times, and having such a positive influence on my life. Scott, I also would not have gotten through my Ph.D. without a coffee machine – best gift EVER! I would also like to acknowledge the support of the Australian Postgraduate Award scholarship and the Multidisciplinary Environment Research Group (MERG) for providing financial support for myself and my innovative research endeavours. Many people have kindly given their time to help improve both the written and intellectual elements of this dissertation – finally, to all co-authors, many thanks! 9 Tall eucalypt forests of Australia: structure, pattern and process SUMMARY The use of observed patterns to infer underlying biotic and physical drivers of eco- evolutionary dynamics is a challenge for population, community and conservation ecology. This is because whilst an understanding of process is required to make useful predictions and test theory, it is static or progressive patterns that are most readily observed in ecological systems, especially at landscape scales. In this thesis, I report on the links between pattern and process, using spatially explicit plot-based information from 48, one hectare tall eucalypt forest communities, spanning the coastal zones of eastern, southern and western Australia. These data are the first to be established across a continental Australian forest system, and were analysed using a combination of point-pattern statistics, likelihood-based inference, structural equation modelling, and simulation modelling. I also developed two new methods for analysing landscape topography and mapping the spatial location of treefalls. Interdisciplinary collaborations, including with astrophysicists, underpinned the novelty of the research. I demonstrated that, at a local scale, climate was a strong predictor of non-eucalypt basal area and density, and eucalypt basal area (but not density), consistently explaining >40% of the deviance in these variables.