The ecology and conservation of bryophytes in Tasmanian wet eucalypt forest Perpetua A. M. Turner B.Sc. (Hons) Submitted in fulfilment of the requirements of the degree of Doctor of Philosophy School of Geography and Environmental Studies University of Tasmania Hobart July 2003 Declaration This thesis contains no material that 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, the thesis contains no material previously published or written by another person, except where due reference is made in the text. Perpetua A.M. Turner (nee Blanks) 11th July 2003 Authority of Access This thesis may be made available forloan. Copying of any part of this thesis is prohibited fortwo years fromthe date this statement was signed; after that time limited copying is permitted in accordance with the Copyright Act 1968. Perpetua A.M. Turner (nee Blanks) 11 th July 2003 11 Abstract The aim of this study was to determine the factors that affect bryophyte richness and species composition in wet eucalypt forest, including old growth and forest disturbed by wildfire or silvicultural practice. Approximately one third of the total bryophyte flora for Tasmania was recorded in old growth mixed forest, with more liverwort than moss species found. Bryophyte species composition was significantly different between groups of sites of forest from the northwest, central and southern areas of the state. Mean annual temperature, altitude, rainfall of the driest month and aspect were most significant in predicting variation in bryophyte species composition. The use of vascular plants as surrogates for the conservation ofbryophyte species was examined. Vascular plant and fem species richness were significant but poor predictors of bryophyte species richness. A minimum set of 31 sites reserved all vascular species and a large percentage (82.9%) ofbryophyte species at least once. Thus, reserves selected using vascular plants are likely to reserve a large proportion of bryophyte species. The reserve sets included more sites of regenerating forest than old growth forest indicating the importance for conservation of multi-aged wet eucalypt forest. Many species preferentially occurred on a substrate type within a particular forest age class. The bryophyte species composition on old growth Nothofagus cunninghamii and Atherosperma moschatum trees were significantly dissimilar to a large number of other substrate/age class groups. Consistent with previous literature, bark type affected species composition. Comparisons ofbryophytes in sites disturbed by wildfire and logging found four moss species occurred more fyequently in logging than wildfire regeneration, whereas six of the seven bryophytes species that occurred more frequently after wildfire than logging disturbance were liverworts. Overall, little difference in bryophyte and vascular species composition was found between logging and wildfire regeneration. When sites were separated into regions, bryophyte species composition differed between logging and wildfire only in the forests of central Tasmania, where Eucalyptus regnans is dominant. 111 Successional stages ofbryophytes species occurrence after disturbance were documented. Species occurring frequently in primary succession did not survive into later successional stages. Many species that established in post-primary successional forest persisted into late successional forest. Liverwort species dominated in late successional forest. The exclusive occurrence of the epiphytic mosses Neckera pennata and Calyptopogon mnioides in regenerating forest is strongly associated with the presence of Pomaderris apetala and Acacia dealbata trees. iv Acknowledgements I am indebted to my supervisors. Jamie Kirkpatrick for his enthusiasm, support, encouragement and wisdom and Emma Pharo, for her knowledge, experience, advice and for keeping me motivated. This PhD and the move to Tasmania would not have been possible without assistance from an APA scholarship. Additional encouragement, advice and support from a number of people also facilitated the move. I thank Mark Burgman for initiating my Honours project based on bryophytes which introduced me to a fascinating new world of plants. I thank the late George A.M. Scott for his wisdom, time, patience and enthusiasm. Without his counsel I would not have seriously considered undertaking a PhD on bryophytes in Tasmania. Mick Brown and John Hickey provided valuable logistic and technical support and taught me a great deal about forest ecology. Their advice, enthusiasm and constant encouragement greatly contributed to the project. I thank Sue Jennings for the tremendous assistance and logistic support while in the field, and for her friendship. Leigh Edwards assited in finding relevant field sites. Keely Ough provided ecological advice and much reference material. Fieldwork would not have been possible without the assistance from Forestry Tasmania staff and Paul Smart, Darren Turner, Emma Pharo, Steve Scott, Steve Davis, Craig Weston, Frank Bishop, Rod Evans, Sandra Hetherington, Bill Tewson and Peter Ladaniwskyi. The patience and unbounded assistance given by Paul Smart is genuinely appreciated. I am indebted to two great friends, Anita Wild and Mark Poll and the 'children', Farmy, Sal and Mush. The support, encouragement, guidance given from Anita and Mark (and licks from the 'children'!) enabled me to achieve many goals. The amazing bryological network was always ready and willing to help at every tum. Advice and assistance were received predominantly with sampling methods and identification of difficult specimens. I am indebted to bryologists in Australia and New Zealand, in particular Dana Bergstrom, Elizabeth Brown, Judith Curnow, Paddy Dalton, Jean Jarman, Neils Klazenga, David Meagher, Pina Milne, Emma Pharo, Helen Ramsay, Rod Seppelt, Jessica Beever, Pat Brownsey, Alan Fife and David Glenny. Assitance was also gratefully received from John J. Engel (Lepidoziaceae and Geocalycaceae), Jan-Peter Frahm (Campylopus), RiclefGrolle (Lepidoziaceae, Lejeuneaceae), Bob Magill, Ray Tangney (Camptochaete) and Kohsaku Yamada (Radula). The assistance and advice provided by David Meagher is greatly appreciated. I am also indebted to staff of the Tasmanian Herbarium, especially Lyn Cave. I am indebted to Leon Barmuta, Lee Belbin, Steve Candy, Richard Little, Peter Minchin and Henrik Wahren for their advice and assitance with some analyses. Thanks to the staff and fellow 'Jamie' students of Geography and Environmental Studies. I thank Moya Kilpatrick and Pauline Harrowby for their assistance and patience. Denis Charlesworth carried out chemical analyses of soil samples. Kate Charlesworth helped find theses, articles and maps. David Somerville assisted with fieldwork equipment. Jamie Kirkpatrick, Emma Pharo, John Hickey, Stephen Bresnehan, Aruta Wild, Ljiljana Sekuljica, Sapphire McMullan-Fisher and Rod Seppelt carried out proofreading. Anita Wild prepared the maps for the chapters. Darren Turner assisted with formatting of this thesis. Thanks especially to Anita and Sapphire for spending time checking the whole thesis. To the Blanks family, I thank you for all your support and understanding. To Nell and Tess whose 'garden landscaping' and waggling tails never failed to bring a smile to my face when things seemed awry. Thanks also to the Turner family for your best wishes and support. And most of all I thank Darren Thank-you for your understanding, love, patience, kindness and commitment, especially during the (many) stressful times. Without you, all this would not have been possible. V Moss Gathering. Theodore Raethke This poem has been removed for copyright reasons Vl Table of Contents Chapter One Introduction ..................................................................................................................... 1 Bryophytes ..................................................................................................................... 1 Bryophytes in forest ecosystems ................................................................................... 1 Wet eucalypt and mixed forest .................................................................................. 1 Fire in wet eucalypt forests ....................................................................................... 2 Silvi cultural practices in wet eucalypt forests .......................................................... .3 The effect of wildfire and logging practices on bryophytes ...................................... 3 Bryophytes in old growth mixed forest.. ................................................................... 5 The importance of substrate in forests ...................................................................... 5 Conservation ofbryophytes in forests ....................................................................... 6 Successional processes in forest ecosystems ............................................................. 8 Thesis aims and structure .............................................................................................. 8 Chapter Two Bryophyte relationships with environment in Tasmanian old growth wet eucalypt forest............................................................................................................................... 11 Abstract ....................................................................................................................... 11 Introduction ................................................................................................................