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Frost Tolerance and Water Relations of Banksia Species in Tasmania Past and Present

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Frost Tolerance and Water Relations of Banksia Species in Tasmania Past and Present FROST TOLERANCE AND WATER RELATIONS OF BANKSIA SPECIES IN TASMANIA PAST AND PRESENT by Julieanne Blake Submitted in fulfilment of the requirements for the degree of Doctor of Philosopy 6e-0 fito,./t June 1997 Declaration This thesis does not contain any material which has been submitted for the fulfilment of any other degree or diploma. To the best of my knowledge and belief, this thesis contains no material which has been published, written or provided by another person, except where due reference is given. J. Blake ACKNOWLEDGMENTS I would like to thank the following people: Dr. Bob Hill for his supervision, Dr. Greg Jordan for his field assistance, discussion and advice on statistical analysis, Dr. Chris Beadle, Dr. Terrence Blake and Dr. Don White for valuable discussion, Kristen Williams and Jane Heath for field assistance, Richard Barnes, Dr. Craig Hardener, Dr. Janet Gorst, Lee Johnson, Peter Bobbii and Gary Haig. I would like to especially thank Mike O'Neill. ABSTRACT Over the past several million years the diversity of Banksia species in Tasmania has declined. On the Tasmanian mainland today there are only two species of Banksia . These are the widespread and morphologically diverse Banksia marginata and Banksia serrata which has a very restricted to the State's northwest. It is not clear when the decline in Banksia began in Tasmania, but Pleistocene sediments in western Tasmania demonstrate a relatively recent extinction for at least some Banksia species. The most recent recorded Banksia extinctions from Tasmania are B. kingii and B. strahanensis. These extinct fossil species are closely related to the extant mainland species B. saxicola/B. canei and B. spinulosa/B. spin ulosa var. cunninghamii respectively. Based on this information it is interesting that B. marginata managed to survive the climatic upheavals of the Quaternary in Tasmania while most other Banksia species were unable to. This thesis investigates whether the extinction of B. kingii and B. strahanensis from Tasmania during the Pleistocene could be linked to a physiological incapacity to tolerate the climatic stresses of the Pleistocene glaciations, i.e. drought and cold. A physiological examination of the drought and cold tolerance of the closest living relatives of the fossil species implies that the frost and drought tolerance of B. kingii and B. strahanensis were no less remarkable than that displayed by a sample of Tasmanian populations of the very successful Banksia marginata. It is therefore concluded that the Pleistocene extinction of B. kingii and B. strahanensis from Tasmania is unlikely to be due to a physiological weakness of drought and cold. This study highlights the physiological mechanisms used Banksia marginata, B. sax/cola, B. cane! and B. spinulosa var. cunninghamiito survive climatic stress. All species were able to significantly increase their bulk elastic modulus in response to drought stress. Banksia marginata and B. spinulosa var. cunninghamii were also able increase their apoplastic water content in response to drought stress. The capacity for osmotic adjustment was not characteristic of any of the species examined. It is interesting that all of the species examined underwent osmotic de-adjustment in response to drought stress. Frost experiments indicate that all of the Banksia examined have reasonable frost tolerance. Although B. spinulosa var. cunninghamii was generally less frost tolerant than the other species, the results were usually significant. The frost results iimply that sensitivity of the fossil species to frost would have unlikely to cause their extinction from Tasmania during the Pleistocene glaciations. Indeed many of the Banksia marginata populations and B. canei and B. sax/cola were able to significantly improve their frost tolerance when exposed to drought stress. The drought and frost experiments in this study successful highlight the similarities of drought and cold stress in their effect on plant physiology. Banksia marginate was examined in more detail than the closest living relatives of the fossil species. This was done to help determine whether its current success in Tasmania is the result of its capacity for physiological plasticity and the genetic diversity of the species. The physiology results indicate that this species is very desiccation tolerant, thus giving it an excellent capacity to deal with drought and cold stress. The physiology results in general also indicated significant differences the drought and frost tolerance among populations of B. marginata. A small isozyme study was performed which suggested there could be a genetic basis to the physiological differences observed among two populations studied along an altitudinal cline. An attempt was also made to determine whether the high degree of morphological variation evident in Tasmanian populations of B. marginate could be positively correlated with physiological variation, which it couldn't. Further work would need to done for this result to be accepted as conclusive. It is likely that the success of Banksia marginata in Tasmania today is related to a combination of genetic diversity of the species, its physiological plasticity and its general hardy nature as evident from its extremely drought tolerant tissues. The extinction of B. kingii and B. strahanensis from Tasmania during the Pleistocene may have resulted during the interglacial periods of the Pleistocene. Glacials are of a much longer duration than interglacials. Interglacials may have been very climatically stressful for the Banksia. Other hypotheses are considered in the thesis. It is also likely that the genetic diversity of B. kingii and B. strahanensis was less than B. marginata during the Pleistocene. This theory is based on the current day morphological variation in the closest living relatives of the fossil species which is considerably less than displayed by B. marginata. In addition, the closest living relatives today have much narrower distributions than B. marginata, hence perhaps having less chance of surviving the climatic stress of the Pleistocene than the more widely dispersed and genetically diverse B. marginata which may have had greater genetic reserves for natural selection to act on. TABLE OF CONTENTS Page CHAPTER 1 Introduction 1 The History of Ban ksia in Tasmania 1 CHAPTER 2 The Role of Plant Physiology in Palaeobotany 18 CHAPTER 3 Details of the Species Studied 22 CHAPTER 4 Water Relations and Frost Methodologies 32 CHAPTER 5 Water Relations Experiments 57 CHAPTER 6 Frost Tolerance Experiments 81 CHAPTER 7 The Cline 99 CHAPTER 8 Genetic Variation Among Two Populations of B. marginata Along an Altitudinal Cline 132 CHAPTER 9 Intrapopulation Variation Within Five Populations of B. marginata 143 CHAPTER 10 General Discussion 156 The Significance of the Physiology Results 156 The Extinction of B. kingii and B. strahanensis from Tasmania 158 REFERENCES 171 APPENDICES Chapter 1 CHAPTER 1. INTRODUCTION The History of Banksia in Tasmania Plant macrofossils and palynological studies provide a window into the past from which information can be drawn about palaeofloras, palaeoclimatic conditions and plant species' extinction. Over the past several million years the diversity of Banksia in Tasmania has undergone decline, e.g. at least seven species with Banksia type leaves were identified from a 35 million year old fossil deposit at Cethana in western Tasmania (Hill et al. 1992). Today, however, apart from the ubiquitous Banksia marginata Cav, there is only one other Banksia species present on the Tasmanian mainland, B. serrata L. f. which is restricted to a very small area in the north-west. Banksia marginata is widely distributed throughout south-eastern Australia today, and is common in Tasmania, South Australia, New South Wales and Victoria (Fig. 1.1). Across its range, B. marginata displays great morphological variation, ranging from a small shrub less than 0.5 m in height, to a tree, taller than 10 m (Taylor and Hopper 1988). Its leaves vary enormously in size and shape (Fig 1.2). Even in the same general area there can be plants with different types of leaves and growth habits (Taylor and Hopper 1988), (Fig. 1.3). Banksia marginata is the most common species of Banksia (Taylor and Hopper 1988). It is tolerant of a wide range of abiotic conditions and even on the fringe of its climatic range it appears to grow well. It occupies a wide range of soil types having a preference for sandy soils, although it is also found on clay loam, peaty loam, rocky soil, quartz, sandstone, limestone and granite (George 1981). It can occur in shrubland, woodland forest, swamps and coastal dunes within a rainfall range of 400-1000 mm per annum (George 1984). In Tasmania, B. marginata ranges from sea level to 1200 m altitude (Salkin 1979). It is common as a tree in western Tasmanian mixed forest and is found on the flood plains of the upper reaches of large rivers such as the Loddon, Franklin and Huon (Salkin 1979). It has also been recorded on the wet, acidic button grass plains of Phillips Lead, Huon Plains, 1 Chapter 1 and Cox Bight (Salkin 1979), all coastal regions, Bruny Island, the Central Plateau and many other areas. It is not clear when the decline in Banksia began in Tasmania but Pleistocene sediments in western Tasmania demonstrate a relatively recent extinction for at least some Banksia species (Jordan and Hill 1991). The most recent recorded Banksia extinctions from Tasmania are B. kingii and B. strahanensis, which have been described by Jordan and Hill (1991) from Pleistocene leaves and infructescences (Figs 1.4, 1.5 and 1.6). These species became extinct during the Pleistocene and are closely related to the extant B. saxicola A.S. George/B. canei J.H. Willis and B. spinulosa var. cunninghamii Sieber ex Reichenbach respectively, which are all currently found on the Australian mainland. In part, this study has sought to explain the extinction of B. kingii and B. strahanensis from Tasmania during the Pleistocene.
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