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I Effects of Factors Associated with the Season of a Fire on Germination Of Effects of factors associated with the season of a fire on germination of species forming soil seedbanks in the fire-prone Hawkesbury Sandstone region of Sydney, Australia. Paul Bengt Thomas B.Sc., The University of Adelaide B. App. Sc. (Hons), The University of Adelaide A thesis submitted for the degree of Doctor of Philosophy University of Western Sydney October, 2004 i Declaration This work contains no material which has been accepted for the award of any other degree or diploma in any university or other 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 has been made in the text I give consent to this copy of my thesis, when deposited in the University Library, being available for loan and photocopying Paul Bengt Thomas October 2004 ii Acknowledgements I would like to thank my supervisors, Charles Morris and Tony Auld for their help and interest throughout this project. This project has benefited from discussions with many people, especially Tony Haigh, and from the technical assistance of Burhan Aramiji. I gratefully acknowledge the help with identification of plant species by staff at the Sydney Botanical Gardens. This work was conducted with the financial assistance of an Australian Postgraduate Award, as part of an ARC SPIRT Grant, and the New South Wales National Parks and Wildlife Service as the industry partner. Seeds were collected under New South Wales National Parks and Wildlife Service Scientific Investigation Licence Number B2063. I would like to thank Chris, Clint, Justin, Matthew and Steve for believing in me. Thank you Stephanie for your love and support during this project. iii Declaration................................................................................................................................. ii Acknowledgements...................................................................................................................iii Abstract ...................................................................................................................................... v Chapter 1. General introduction.................................................................................................1 Chapter 2. Effects of fire-related germination cues ................................................................. 10 2.1 Introduction.................................................................................................................... 10 2.2 Methods.......................................................................................................................... 18 2.3 Results............................................................................................................................ 30 2.4 Discussion ...................................................................................................................... 56 Chapter 3. Effects of pre- and post-fire temperature................................................................ 68 3.1 Introduction.................................................................................................................... 68 3.2 Methods.......................................................................................................................... 79 3.3 Results.......................................................................................................................... 101 3.4 Discussion .................................................................................................................... 159 Chapter 4. Effects of pre-fire hydration status....................................................................... 170 4.1 Introduction.................................................................................................................. 170 4.2 Methods........................................................................................................................ 176 4.3 Results.......................................................................................................................... 185 4.4 Discussion .................................................................................................................... 205 Chapter 5. Effects of post-fire water...................................................................................... 211 5.1 Introduction.................................................................................................................. 211 5.2 Methods........................................................................................................................ 221 5.3 Results.......................................................................................................................... 236 5.4 Discussion .................................................................................................................... 272 Chapter 6. Effects of fire on germination of aged seed.......................................................... 286 6.1 Introduction.................................................................................................................. 286 6.2 Methods........................................................................................................................ 291 6.3 Results.......................................................................................................................... 293 6.4 Discussion .................................................................................................................... 306 Chapter 7. General discussion................................................................................................ 309 References:............................................................................................................................. 314 Appendix 1......................................................................................................................... 345 iv Abstract Fire is a recurrent disturbance that removes above ground vegetation in many locations throughout the world, including the Sydney region. Many species in fire-prone locations, and most species in the Sydney region, form soil seedbanks and regenerate through post-fire germination. However, a germination response is determined by the fire regime acting as a selective pressure over a sufficient period of time, rather than a single fire. The components of the fire-regime are intensity, season, type and frequency. The natural fire- regime is dominated by warm-season fire, but management burning is conducted in cooler seasons. Cool season burning produces lower levels of germination than warm season fires in a number of locations with Mediterranean-type climate, but the effects of cool season burning on species composition in the relatively aseasonal Sydney region is unknown. An experimental approach was adopted to address this lack of knowledge. Fire can be simulated using heat shock and smoke (fire cues), and the seasonal factors of temperature and water- availability can be reproduced in the laboratory. I have investigated the effect of various combinations of heat shock and smoke, of various pre-and post-fire cue temperatures, of pre- fire cue hydration status, of various post-fire cue water availabilities, and of accelerated aging before application of fire cues on germination of a number of species forming soil seedbanks in the Sydney region. A degree of primary dormancy was overcome in most species by the combination of heat shock and smoke in the current investigation. Fire intensity is expected to influence germination, as germination of most species was increased by the combination of heat shock and smoke within a narrow heat shock range. Consequently, both post-fire germination and a residual soil seedbank are expected, with the residual soil seedbank providing a buffer against short fire-return intervals. Season of fire is also expected to influence species composition because both temperature and water availability affected germination in experiments. Ambient temperature affected germination directly, and possibly through secondary dormancy. Germination of a similar proportion of species was favoured by the ambient soil temperature during cooler seasons, as was favoured by warmer season temperature, as was favoured by neither temperature. Generally the effect of temperature was of low magnitude. When temperature had a major effect, then the fire-related germination cues extended the range of ambient temperature over which germination could occur, but was sub-compensatory for optimal temperature. Thus, a similar proportion of species would be expected to germinate over v different season of fire, but species composition would be expected to change with repeated single-season fire. The fire-related germination cues were promotive for a finite duration of post- treatment incubation, thus species that are not favoured by the ambient temperature at the time of the fire are likely to not germinate, rather than merely delay germination until a change of season. When water was intermittently available prior to continuous hydration, then the influence of temperature was strengthened in the single species investigated. The fire-related germination cues followed by dry storage prior to hydration probably caused secondary dormancy. This contrasts with species from Mediterranean-type climates (where the natural fire season and season with adequate rainfall for seedling recruitment are separated
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