The Taxonomy and Ecology of Wood Decay Fungi in Eucalyptus Obliqua Trees and Logs in the Wet Sclerophyll Forests of Southern Tasmania
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The taxonomy and ecology of wood decay fungi in Eucalyptus obliqua trees and logs in the wet sclerophyll forests of southern Tasmania by Anna J. M. Hopkins B.Sc. (Hons.) School of Agricultural Science, University of Tasmania Cooperative Research Centre for Forestry A research thesis submitted in fulfilment of the requirements for the Degree of Doctor of Philosophy January, 2007 Declarations This thesis contains no material which has been accepted for a degree or diploma in any university or other institution. To the best of my knowledge, this thesis contains no material previously published or written by another person, except where due acknowledgment is made in the text. Anna J. M. Hopkins This thesis may be made available for loan and limited copying in accordance with the Copyright Act of 1968. Anna J. M. Hopkins ii Abstract The wet sclerophyll forests in southern Tasmania are dominated by Eucalyptus obliqua and are managed on a notional silvicultural rotation length of 80 to 100 years. Over time, this will lead to a simplified stand structure with a truncated forest age and thus reduce the proportion of coarse woody debris (CWD), such as old living trees and large diameter logs, within the production forest landscape. Course woody debris is regarded as a critical habitat for biodiversity management in forest ecosystems. Fungi, as one of the most important wood decay agents, are key to understanding and managing biodiversity associated with decaying wood. In Australia, wood-inhabiting fungi are poorly known and the biodiversity associated with CWD has not been well studied. This thesis describes two studies that were undertaken to examine the importance of CWD as habitat for wood-inhabiting fungi in the wet sclerophyll forests of Tasmania. Study one examined the effect of changing tree age on the fungal species richness, fungal community composition and rotten wood associations within the tree. Six living E. obliqua trees in each of three age-classes (69, 105 and >150 years old) were felled. Each tree was cross-cut at nine standard sampling points and the decay profile was mapped. Fungi were isolated from rotten wood at each sampling point, and from control samples of clear heartwood and sapwood. Samples of each rotten wood type were collected and classified, based on their colour and texture. Wood-inhabiting fungi in Australia are not well known, making the identification of fungal cultures problematic. In study one, cultures of wood-inhabiting fungi were grouped into putative species groups and identified using both morphological techniques and polymerase chain reaction (PCR) and sequencing of the internal transcribed spacer (ITS) region of the ribosomal DNA. Ninety-one species of wood-inhabiting fungi were isolated from the 18 trees examined in study one. Eight different rotten wood types were identified and studied. The community composition of wood-inhabiting fungi in trees greater than 150 years old (the oldest age-class examined) was very different compared with those found in the younger two age-classes; more than half of all species were only iii found in these older trees. In addition, trees in the oldest age-class contained greater volumes and proportions of rotten wood habitat. The second study examined the wood-inhabiting fungi that had been isolated in a previous study by Z-Q Yuan (University of Tasmania) from large (>85 cm) and small (30-60 cm) diameter E. obliqua logs in mature, unlogged forests and 20-30 year-old logged forests that were regenerating after clearfelling. The previously described morphospecies were tested using PCR-sequencing of the ITS region of ribosomal DNA and a consensus final species groupings was obtained. This study determined that a total of 60 species of wood-decay fungi had been commonly isolated from the 36 logs examined. Significant differences in fungal community structure were found between mature forests and regenerating forests. Some differences in fungal species richness and community composition were also found between logs of different sizes. These studies are among the first to examine wood-inhabiting fungi in mature E. obliqua trees and logs in Tasmania. The ecological information obtained from this research will assist in the development and deployment of strategies for the management of mature living trees and logs in wet eucalypt forests in Tasmania. This research suggests there is a need for forest managers to consider instigating measures that allow for some trees in the production forest landscape to live long enough to develop decayed wood habitat. This will provide important habitat for fungi as both trees and large diameter logs, sustaining an important component of forest biodiversity. iv Acknowledgements During the course of my study I was supported by an Australian Postgraduate Award and a CRC for Sustainable Production Forestry scholarship. Significant in-kind support was also provided by Forestry Tasmania. Operating funds were supplemented by a Warra Research Grant and a grant from the Holsworth Wildlife Research Fund. I sincerely thank my supervisors Dr Caroline Mohammed, Dr Simon Grove and Dr Tim Wardlaw for their wisdom and encouragement throughout this project. I am most grateful for the opportunity to work with them in such an exciting field of endeavour. Dr Morag Glen provided especially helpful guidance in the molecular analyses of my fungi and has been a wonderful support throughout the thesis writing process. Kate Harrison assisted with the initial field work for Study 1 and has been a wonderful support during the analysis and writing process, providing much helpful feedback and discussion. Malcolm Hall, Andrew Muirhead, Leigh Edwards, Jill Butterworth, William Young and Danielle Wiseman assisted with the field work for Study 1. Jack Simpson, formerly of State Forests NSW, taught me all I know about describing wood decay fungi in culture and was instrumental in beginning the morphological descriptions of cultures in Study 1. Dr Zi-Qing Yuan and Dr Marie Yee did the field work for Study 2. In addition, all the isolations, culturing and fungal morphology for study 2 were done by Dr Zi-Qing Yuan. Genevieve Gates and Dr David Ratkowsky collected and identified the fruitbodies used for the local Tasmanian reference collection. Dr Karina Potter and Trang Tranh provided the sequences for the Clayton Collection and occasionally assisted with molecular work. v The support of my fellow postgraduate students Kate Harrison, Lizz Pietrzykowski, Anna Smith and Danielle Wiseman has been invaluable; as has the regular support and feedback from other members of the forest health research group, especially Karina Potter, Karen Barry and Marie Yee. The completion of this thesis would not have been possible without the wonderful support from my friends and family. In particular I would like to thank Lucy Hopkins, Angas Hopkins, Lekkie Hopkins and Anne Nicholls for being so patient, interested and understanding. vi Communication of results from this thesis (to date) Journal Articles (Refereed) Hopkins, AJM , Harrison, KS, Grove, SJ, Wardlaw, TJ, Mohammed, CL. (2005). Wood decay fungi and beetle assemblages associated with living Eucalyptus obliqua trees: early results from studies at the Warra LTER Site, Tasmania. Tasforests 16:111-126. Conference Proceedings Harrison, KS, Hopkins, AJM , Grove, SJ, Wardlaw, TJ and Mohammed, CL. (2004). Tree Age, Invertebrates and Fungi. Poster presentation. 22 nd International Congress of Entomology 2004. 16-21 August 2004, Brisbane, Queensland. Harrison, KS, Hopkins, AJM , Grove, SJ and Mohammed, CL. (2004). Saproxylic invertebrates in living trees in Tasmania’s southern forests. Oral presentation. Ecological Society of Australia Ecology 2004 Conference. 7-10 December, Adelaide, South Australia. Harrison, KS, Hopkins, AJM , Grove, SJ and Mohammed, CL. (2003). Living Trees - what lives here? Saproxylic invertebrates and wood decay fungi in living Eucalyptus obliqua in Tasmania's wet sclerophyll forests. Poster presentation. 34th Australian Entomological Society and 6th Invertebrate Biodiversity Conference. 29 September - 3 October, Hobart, Tasmania. Harrison, KS, Hopkins, AJM , Grove, SJ and Mohammed, CL. (2003). Artificial wounding of living Eucalyptus obliqua in Tasmania's wet sclerophyll forests. Poster presentation. Ecological Society of Australia Ecology 2003 Conference. 8-10 December, Armidale, New South Wales. Hopkins, AJM (2006). Wood-inhabiting fungi in Eucalyptus obliqua logs: examining the effect of log size and forest type on fungal community composition. Oral presentation. The Joint Conference of the NZ Ecological Society and the Ecological Society of Australia. 28 August-1 September, Wellington, New Zealand. Hopkins, AJM (2005). Wood decay fungi and saproxylic invertebrates in Tasmania’s southern forests. Invited oral presentation. Third National Fungimap Conference. 28 April – 3 May 2005, Tasmania. vii Hopkins, AJM (2003). Wood decay fungi and saproxylic invertebrates in Tasmania’s southern forests. Oral presentation. 6 th Australasian Mycological Society Conference & 150 years: National Herbarium of Victoria & Plant Systematics in Australia Anniversary Conference. 29 September-23 October, 2003, Melbourne, Victoria. Hopkins, AJM , Harrison, KS, Grove, SJ, Wardlaw, TJ and Mohammed, CL. (2006). Decayed wood, wood-inhabiting fungi and saproxylic beetles in living trees in southern Tasmania. Poster presentation. Eighth International Mycological Congress. 20-25 August, Brisbane, Australia. Hopkins, AJM , Harrison, KS,