Recruitment of Banksia Spp. in an Anthropogenically Disturbed Mediterranean Climate Type Woodland in Western Australia
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Recruitment of Banksia spp. in an anthropogenically disturbed mediterranean climate type woodland in Western Australia This thesis is presented for the degree of Doctor of Philosophy of Murdoch University 2011 Submitted by Roberto Crosti Bachelor with Honours of Natural Science (University “La Sapienza” Rome Italy) 1 Table of Content DECLARATION 4 ACKNOWLEDGEMENTS 5 LIST OF FIGURES AND TABLES 6 ABSTRACT 14 GENERAL SUMMARY OF THE DIFFERENT CHAPTERS OF THE THESIS 17 1 CHAPTER 1: BANKSIA WOODLANDS ON THE SWAN COASTAL PLAIN 24 1.1 INTRODUCTION 25 1.1.1 CHARACTERISTICS OF BANKSIA WOODLANDS 26 1.1.2 FLORA OF BANKSIA WOODLANDS 27 1.1.3 THE SOIL SYSTEM 28 1.2 VEGETATION CHANGES IN BANKSIA WOODLANDS 30 1.2.1 IMPACTS WITHIN BANKSIA WOODLAND 31 1.2.2 INTER -RELATED PERTURBATIONS 37 1.3 THE LOCATIONS OF THE STUDY 37 1.3.1 KINGS PARK BUSHLAND (KP) 37 1.3.2 THE OTHER LOCATIONS AT WHICH INVESTIGATIONS WERE UNDERTAKEN 41 1.4 THE INVESTIGATED SPECIES 44 2 CHAPTER 2 THE VEGETATION CHANGES STUDY 46 2.1 INTRODUCTION 47 2.2 AIM 51 2.3 MATERIALS AND METHODS 51 2.3.1 THE 1939 MAPPING 52 2.3.2 THE 1999 RE -SURVEYING 52 2.4 RESULTS 56 2.5 DISCUSSION 62 3 CHAPTER 3: SEED PRODUCTION AND DISPERSAL 70 3.1 INTRODUCTION 71 3.2 AIM 73 3.3 MATERIALS AND METHODS 73 3.3.1 TAGGED INFLORESCENCES 74 3.3.2 SEED TRAPS 75 3.3.3 QUADRATS USED TO ESTIMATE POST -FIRE SEED FALL 77 3.4 RESULTS 79 3.4.1 TAGGED INFLORESCENCES 79 3.4.2 SEED TRAPS 89 3.4.3 QUADRATS USED TO ESTIMATE POST -FIRE SEED FALL 96 2 3.5 DISCUSSION 98 4 CHAPTER 4: SEED PREDATION 102 4.1 INTRODUCTION 103 4.2 AIM 107 4.3 PRE -DISPERSAL SEED PREDATION 107 4.3.1 THE STUDY ON PRE -DISPERSAL SEED -PREDATION 109 4.3.2 MATERIAL AND METHODS 109 4.3.3 RESULTS 112 4.4 POST -DISPERSAL SEED PREDATION 118 4.4.1 THE STUDY OF POST -DISPERSAL PREDATION 120 4.4.2 MATERIAL AND METHODS 121 4.4.3 RESULTS 128 4.5 GENERAL DISCUSSION 150 4.5.1 PRE -DISPERSAL PREDATION 150 4.5.2 POST -DISPERSAL PREDATION 153 5 CHAPTER 5: SEED GERMINATION AND SEEDLING SURVIVAL 160 5.1 INTRODUCTION 161 5.2 AIM 163 5.3 GENERAL MATERIALS AND METHODS 163 5.3.1 SURVEYS AND EXPERIMENTS 163 5.3.2 SEEDS 164 5.3.3 GREENHOUSE 165 5.3.4 TYPES OF SUBSTRATA 166 5.3.5 STATISTICAL ANALYSIS 168 5.4 AIM , METHODS , RESULTS , DISCUSSION OF SINGLE SURVEYS AND EXPERIMENTS 169 5.4.1 A: TRANSECTS TO QUANTIFY NATURAL GERMINATION OF BANKSIA IN THE BUSHLAND IN DIFFERENT TYPES OF ENVIRONMENT (INCLUDING SANDY TRACKS ) 169 5.4.2 B: SEED MANIPULATION ON DIFFERENT TYPES OF SUBSTRATA (IN KINGS PARK BUSHLAND WITH THE DOMINANT SPECIES ) 181 5.4.3 C & D: SEEDS TESTED FOR GERMINATION IN NATURAL HABITAT CONDITIONS WITH BANKSIA SPP . (SEEDS PLACED IN AUTUMN AND IN EARLY SUMMER IN THE SOIL SEED BANK ) 188 5.4.4 E: BANKSIA SEEDS SOWN IN THE BUSHLAND IN DIFFERENT TYPES OF ENVIRONMENT IN A 1X1 METRE QUADRAT SCHEME 208 5.4.5 F: EXPERIMENT IN THE GLASSHOUSE , INVESTIGATING GERMINATION OF BANKSIA SPP , ON DIFFERENT TYPES OF SUBSTRATA 215 5.4.6 G: EXPERIMENT IN THE TUNNEL HOUSE WITH ALL SPECIES ON DIFFERENT SUBSTRATA AS IN THE B FIELD EXPERIMENT 221 5.5 GENERAL DISCUSSION 227 6 CHAPTER 6: CONCLUSION 232 6.1 IMPLICATIONS FOR CONSERVATION AND MANAGEMENT 241 REFERENCES 244 APPENDICES 267 3 DECLARATION I hereby declare that this submission is my own work and that, to the best of my knowledge and belief, it contains no material previously published or written by another person nor material which has been accepted for the award of any other degree or diploma of the university or other institute of higher learning, except where due acknowledgment has been made in the text. 4 ACKNOWLEDGEMENTS This thesis took a lot of time to be completed and was possible thanks to the support of many people. I am really grateful to my main supervisor Phil Ladd for his combined expertise, experience, patience and his “trans-hemisphere” frequent flights. Phil taught me a great deal about research and helped throughout the project, in the field, in laboratory and with thesis production. I was mainly based at the Science Directorate at the Botanic Garden and Parks Authority (BGPA) and I’m really grateful to Kingsley Dixon who supervised me in the research in the Kings Park bushland and in the experiment undertaken in the greenhouse facilities. Thanks are also due to the staff and students of the Murdoch University and of the Botanic Gardens and Parks Authority for assistance and use of facilities. In particular I really would like to thank: Bob Dixon, Deanna Rokich, Siegy Krauss, Keran Keys, Anlie Tieu, Colin Yates, Matt Barrett, David Merritt, Russell Barrett, Katinka Ruthrof, Judy Fisher, Grant Wells, Ray Wills, Patrick Courtney, Michelle Carey, Andrew Batty and Tissa Senaratna Also Erika Wikus and Sandro Pignatti that always encouraged my research I need also to thank my mum and my sister for continuously asking...” when are we all going to Australia for the thesis ceremony? ” Finally to my beloved wife Antonella who helped in the field, supported me writing the thesis and always encouraged me to finish it. With Isabel and Nicola we are a beautiful family and now we will have more time to spend together. 5 LIST OF FIGURES AND TABLES Chapter 1 Fig. 1.1 Distribution and extent of Banksia woodlands on the Swan Coastal Plain, in the southwest of Western Australia (modified from Beard 1989). Fig. 1.2 Soil systems of the Perth region (McArthur and Bettenay 1974). Fig. 1.3 The Australian continent and the Swan Coastal Plain. In the rectangular the Perth region where the investigated sites are located. KP= Kings Park; BP= Bold Park; Mu= Murdoch bushland; Rid= Ridges State Forest; Jand= Jandakot Regional Park. Chapter 2 Fig. 2.1 Perth, Western Australia and location of the study plots in Kings Park. Fig. 2.2 Thermopluviogram data taken from the nearest weather station to the study area, averages of temperature and precipitation from the 20 year periods before 1939 and 1999. The temperature bar chart and precipitation lines show this is a typical mediterranean climatic regime. Fig. 2.3 Comparison between plant density of the selected study species (hectare -1), in 1939 and 1999; mean ± 1SE based on 126 transects. Fig. 2.4 Comparison between plant relative frequency of the selected study species, in 1939 and 1999. Fig. 2.5 Structure of the Kings Park bushland: above 1939 and below 1999. Fig. 2.6 Non-metric multidimensional scaling ordination of 1939 and 1999 plots. NMDS in two dimensions with Bray-Curtis distance, function Stress = 0.07 Table 2.1 General attributes of the plots where transects were located in 1939 and resurveyed in 1999 in Kings Park. S=surveyed plots, N=number of transects, F=number of extensive wildfire in the 60 years period Table 2.2. Climate in the study area, for the 20 year periods prior to 1939 and 1999; Mx=mean maximum air temperature, Mn= mean minimum air temperature, Dm=absolute mean, P=Precipitation, S=Bright sunshine. Table 2.3. Mean values of trees (hectare -1) for transects within each plot and statistical significance, using t-test and sign test, based on paired comparison, between 1939 and 1999, of 126 transects; **=P<0.01, *** =P<0.001. Table 2.4. Mean values of shrubs (hectare -1) for transects within each plot and statistical significance, using t-test and sign test, based on paired comparison, between 1939 and 1999, of 126 transects; **=P<0.01, *** =P<0.001; ns =not significant. Chapter 3 6 Fig. 3.1 B. attenuata and B. menziesii closed follicles. Mean percentage of the different sites in the four locations. Fig. 3.2 B. attenuata closed follicles . Mean percentage of the different sites for each of the four locations. Fig. 3.3 B. menziesii close follicles . Percentage of the different sites for each of the four locations. Fig. 3.4 B. attenuate mean of open, closed, predated/damaged follicles for “cone”. Fig. 3.5 B. attenuata mean percentage of developed and undeveloped “cones”, on the tree or fallen from the tree. Fig. 3.6 B. menziesii mean number of open, close, predated/damaged follicles for “cone”. Fig. 3.7 B. menziesii mean percentage of developed and undeveloped “cones”, on the tree or fallen from the tree. Fig. 3.8 B. attenuata developed fallen infructescences. Percentage of the different sites in the four locations. Fig. 3.9 B. attenuata developed fallen infructescences. Average percentage of the different sites for each of the four locations. Fig. 3.10 B. menziesii developed fallen infructescences. Percentage of the different sites in the four locations. Fig. 3.11 B. menziesii developed fallen infructescences. Average percentage of the different sites for each of the four locations. Fig. 3.12 B. attenuata undeveloped fallen infructescences. Percentage of the different sites in the four locations. Fig. 3.13 B. attenuata undeveloped fallen infructescences. Average percentage of the different sites for each of the four locations. Fig. 3.14 B. menziesii undeveloped fallen infructescences. Percentage of the different sites in the four locations. Fig. 3.15 B. menziesii undeveloped fallen infructescences. Average percentage of the different sites for each of the four locations. Fig. 3.16 B. attenuata and B. menziesii mean number of seeds m -2 y -1.