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1999 A holocene vegetation history of the Flinders Ranges South Australia: evidence from Leporillus spp. (stick-nest rat) middens Lynne McCarthy University of Wollongong

Recommended Citation McCarthy, Lynne, A holocene vegetation history of the Flinders Ranges South Australia: evidence from Leporillus spp. (stick-nest rat) middens, Doctor of Philosophy thesis, School of Geosciences, University of Wollongong, 1999. http://ro.uow.edu.au/theses/1962

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A HOLOCENE VEGETATION HISTORY OF THE FLINDERS RANGES SOUTH AUSTRALIA: EVIDENCE FROM LEPORILLUS SPP. (STICK-NEST RAT) MIDDENS

A thesis submitted in fulfilment of the requirements for the award of the degree

DOCTOR OF PHILOSOPHY

from

UNIVERSITY OF WOLLONGONG

by

LYNNE MCCARTHY B.Env.Sc. BSc (Hons.)

SCHOOL OF GEOSCIENCES 1999 This work has not been submitted for a higher degree at any other University or Institution and, unless acknowledged, is my own work.

Lynne McCarthy i

ABSTRACT

Palaeoecological records for semi-arid and arid environments of Australia are limited due to poor preservation of material in this environmental setting. As a consequence, a Holocene vegetation and climatic record for a large part of the continent is incomplete. Leporillus spp. (stick-nest rat) middens provide a wealth of palaeoecological information for Holocene environments in areas where such records are rare. Eighteen middens from three key sites in the Flinders Ranges (Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge), were investigated in this project to provide a thorough spatial and temporal coverage of palaeoecological sites. Issues of midden taphonomy, temporal resolution of pollen and macrofossil evidence, refining the interpretation of palaeoecological records from middens, and reconstructing palaeovegetation and climates during the Holocene, are dominant themes of this research.

Modern pollen rain at study sites was investigated to provide the foundation for interpretation of palaeoecological evidence from the stick-nest rat middens. A study of the regional modern pollen rain along a west-east transect in the central and northern Flinders Ranges reflected the high spatial variability in vegetation communities from semi-arid rocky upland environments. Local environmental conditions at individual midden sites (aspect and degree of exposure at cave sites and local vegetation cover) were significant factors in determining the composition of the modern pollen rain at individual midden sites .

Accelerator Mass Spectrometry (AMS) radiocarbon dating was used to examine the taphonomy of middens and refine the temporal resolution of these deposits. The majority of middens were found to have been deposited rapidly. Temporal resolution of middens can be complex, however in most instances, there were contemporaneous ages of different components in the middens (leaves, pollen concentrates and faecal pellets). AMS dating provided reliable ages for individual middens.

Vegetation taxa were better represented by pollen rather than macrofossils in the midden assemblages as evidenced by MacrofossihPollen Index values. Macrofossils are regarded as a secondary line of evidence in these midden records. They can verify the local occurrence of a taxon, whose source (local or regional) may not be distinguished solely in the pollen record.

Pollen and macrofossil records suggest that woodland and shrubland communities with an understorey of herbaceous taxa and grasses were dominant around 7 000-5 000 yrs B.P in the northern ranges, and shrublands with an understorey of herbaceous taxa and chenopods were dominant in the central ranges. This is indicative of wetter and warmer ii

conditions than present during this part of the Holocene. Shrubland communities declined in the central ranges while persisting in the north from 4 000-2 000 yrs B.P, to be replaced by chenopod shrublands with a less diverse herbaceous component in the understorey with increasing aridity into the Late Holocene. Chenopod shrublands continued to increase from 1 000 yrs B.P to the present in the central ranges. Midden records have provided evidence for a shift between transitional shrubland communities to chenopod shrublands in the central ranges. To the north, woodland and shrublands remained throughout the Holocene. Topographically buffered vegetation in the northern ranges is more resilient than central ranges vegetation to climate change.

Change in vegetation communities was more visible in the central ranges, as a result of different environmental conditions more sensitive to changes in effective precipitation. This highlights the complexity of factors that affect plant distributions in semi-arid environments. Underlying the climatically driven response is the role of biogeographic parameters that influence the diversity and structure of different vegetation communities. In sheltered rugged topography, in the northern ranges, plant communities are more stable as evidenced by the occurrence of relict species and maintenance of woodland and shrubland communities throughout the Holocene even as conditions were becoming increasingly arid. The same climatic scenario of increasing aridity in the central ranges resulted in less stable vegetation communities that responded to cooler and drier conditions by shifting from dominantly shrublands to an extensive cover of chenopod shrublands. Present spatial variability in the vegetation being a feature of the last 1 000 yrs (and possibly longer in the central ranges), compared with more homogeneous conditions across the ranges from 7 000-5 000 yrs B.P, is certainly consistent with the argument that climatic parameters such as ENSO have become more variable during the Late Holocene. iii

ACKNOWLEDGEMENTS

Many thanks to Lesley Head for constant support, encouragement, patience and guidance during different stages of the PhD. Thankyou for the generous amount of time invested into reading draft chapters, useful discussions, and providing thoughtful and constructive comments about this work, especially during the writing stage.

Thankyou to a team of people who provided invaluable assistance and expertise in their contributions to different aspects of this thesis. Graham Medlin for his enthusiasm and generosity in providing assistance with bone macrofossil identifications, locations of middens and helpful discussions. Helen Vonow, Martin O'Leary and staff at the Adelaide State Herbarium for plant and macrofossil identifications. Professor Michael Tyler for frog bone identifications. Greg Martin for assistance in finding middens. Trevor Nasmith and Eric Dahl from the South Australian Parks and Wildlife for assistance in the field. Botanists from the Department of NSW Agriculture Seed Laboratory for identifications of fossil seeds. BETA Analytic and ANSTO for AMS radiocarbon dates. Sue Pritchard, David Martin and Richard Miller for drafting wonderful diagrams and maps. Robert McCarthy and Lesley Head for proof reading final drafts of the thesis. Judy McCarthy, Jan McCarthy, Jenny Atchison, Vanessa Allen, Terry Lachlan, Jeannette Mangan and Lesley Head for help and plenty of laughs during the hunt for amberat, pollen traps and vegetation surveys during field trips to the Flinders Ranges. The study was funded by the ARC (Grant no. A19530447), AINSE (Grant no. 94/188 and 97/116R) and the Quaternary Environments Research Centre, University of Wollongong.

I am grateful to Julio Betancourt, Tom Van Devender and families, for their generosity and hospitality during my visit to Tucson. The opportunity to work with, and exchange ideas and information with such respected researchers in the field of midden analysis was valuable.

Thankyou to great friends Jenny, Maria, Laurie, Filiz, David, Gordon and John who have kept me sane and have been a source of strength and happiness, especially during the final stages of putting this thesis together. Special thanks to Brendan for love and support right from the beginning and for always seeing great things ahead for me.

Thank you to Dad, Judy, Jan, Robert, Gerry and Dawn for always believing that I could do this and finish it! IV

TABLE OF CONTENTS

Abstract i Acknowledgements iii Table of Contents iv List of Plates ix List of Figures x List of Tables xv

CHAPTER 1: INTRODUCTION: THESIS AND AIMS 1.1 Palaeoecology of the Semi-Arid Zone: the issues, questions and problems 1 1.2 The Evidence 1 1.3 The Thesis: Methodological and Taphonomic Issues 3 1.4 Thesis Objectives 3 1.5 Chapter Outline 5

CHAPTER 2: THE RESEARCH CONTEXT 2.1 Introduction 7 2.2 Midden Analysis 7 2.2.1 Neotoma Midden Analysis: the beginnings 8 2.2.2 Contributions of Neotoma Midden Studies to the Palaeoecology of Desert Areas in the USA 9 2.2.3 Midden Studies in South Africa and the Middle East 11 2.2.4 Methodological and Taphonomic Issues in Midden Analysis 13 2.2.5 Australian Midden Studies 18 2.3 Holocene Palaeoenvironments of Australia 22 2.3.1 Overview of Holocene Climate 23 2.3.2 Lake Level and Pollen Records 24 2.3.3 Dune and Fluvial Records 27 2.3.4 Evidence for Seasonality in Holocene Climates 27 2.3.5 Contradictions Between Holocene Climate Reconstructions 28 2.4 Contribution of Leporillus spp. Midden Records 30 2.5 Conclusion 31

CHAPTER 3: THE FLINDERS RANGES: A REGIONAL PERSPECTIVE 3.1 Introduction 33 3.2 The Flinders Ranges 33 3.2.1 Geological Setting 33 V

3.2.2 Climate 35 3.3 The Semi-Arid Environment: Factors Influencing Vegetation Communities 37 3.3.1 Physical Environment 37 3.3.2 Influence of Rainfall 37 3.3.3 The Role of Disturbance 38 3.4 Semi-Arid Vegetation Communities: Sub-Continental Context 41 3.4.1 Tree and Shrub Species 41 3.4.2 Chenopod Shrublands 42 3.4.3 Semi-Arid Woodlands 42 3.4.4 Arid and Semi-Arid Low Woodlands 43 3.4.5 Acacia Shrublands 43 3.4.6 Grasslands 43 3.5 Vegetation Communities in the Northern and Central Flinders Ranges: the Regional Context 44 3.5.1 Riverine and Dry Semi-Arid Woodlands 44 3.5.2 Low Open Woodland 45 3.5.3 Mallee Shrublands 45 3.5.4 Tall Shrublands 47 3.5.5 Chenopod Shrublands 47 3.5.6 Tussock Grasslands 50 3.6 Sensitivity of the Flinders Ranges: a refuge 50 3.7 Regional Setting and Midden Study Sites: implications for this thesis 51

CHAPTER 4: METHODOLOGIES: STRATEGIES AND TECHNIQUES FOR MIDDEN ANALYSIS 4.1 Introduction 52 4.2 Overview of Research Strategy 52 4.3 Arkaroola-Mount Painter Study Site 53 4.4 Mount Chambers Gorge Study Site 53 4.5 Brachina Gorge Study Site 54 4.6 Midden Survey and Sampling 56 4.7 Field Sampling of Modern Vegetation Communities 57 4.8 Pollen Trap Studies 58 4.9 Laboratory Methods 61 4.9.1 Stick-nest Rat Midden Macrofossil Recovery 61 4.9.2 Identification of Macrofossils 62 4.9.3 Pollen Recovery from Traps 64 4.9.4 Pollen Reference Material 64 4.10 Pollen Counting and Identification 64 vi

4.11 Accelerator Mass Spectrometry (AMS) Radiocarbon Dating of Macrofossils and Pollen 65 4.11.1 Pollen Concentration for AMS Radiocarbon Dating 66 4.12 Statistical Analysis 67 4.13 Conclusion 69

CHAPTER 5: INVESTIGATION OF MODERN POLLEN RAIN IN THE FLINDERS RANGES 5.1 Introduction 70 5.2 Understanding Modern Pollen Rain 70 5.2.1 Pollen production, dispersal and deposition 70 5.2.2 Finding a Modern Analogue 71 5.2.3 Representativeness of Pollen Taxa 71 5.2.4 Sampling Sites and Interpretation of Pollen Assemblages 72 5.3 Pollen Traps and Vegetation Data 74 5.3.1 Arkaroola Pollen Trap Transect and Vegetation Data 74 5.3.2 Mount Chambers Gorge Pollen Trap Transect and Vegetation Data 83 5.3.3 Brachina Gorge Pollen Trap Transect and Vegetation Data 89 5.4 Cluster Analysis of Transect Pollen Trap Study 95 5.4.1 Indicator Pollen Taxa 102 5.5 Pollen Trap Studies at the Midden Cave Sites 108 5.5.1 Arkaroola Cave Traps and Vegetation Surveys 108 5.5.2 Mount Chambers Gorge Cave Traps and Vegetation Surveys 136 5.5.3 Brachina Gorge Cave Traps and Vegetation Surveys 144 5.6 Modern Pollen Spectra at Midden Cave Sites 160 5.7 Comparison Between Transect and Cave Trap Studies 161 5.8 Conclusions 163

CHAPTER 6: STICK-NEST RAT MIDDEN PALAEOECOLOGICAL RECORDS FROM THE NORTHERN FLINDERS RANGES: ARKAROOLA-MOUNT PAINTER SANCTUARY 6.1 Introduction 165 6.2 Arkaroola Middens 165 6.2.1 Waterfall 1 (WF1) Midden 165 6.2.2 North Well Creek 1 (NWCK1) Midden 176 6.2.3 North Well Creek 2 (NWCK2) Midden 179 6.2.4 Radium Creek 1 (RC1) Midden 180 6.2.5 Radium Creek 2 (RC2) Midden 180 6.2.6 Oppaminda Track (OT1) Midden 181 Vll

6.2.7 Radium Creek 3 (RC3 TOP) and (RC3 BASE) Midden 182 6.2.8 Haematite Hill (HH1 TOP) and (HH1 BASE) Midden 183 6.2.9 Arkaroola 1 (ARK1 TOP), (ARK1 MIDDLE) and (ARK1 BASE) Midden 183 6.3 Conclusion 187

CHAPTER 7: STICK-NEST RAT MIDDEN PALAEOECOLOGICAL RECORDS FROM THE CENTRAL FLINDERS RANGES: MOUNT CHAMBERS GORGE AND BRACHINA GORGE 7.1 Introduction 188 7.2 Mount Chambers Gorge Middens 188 7.2.1 Mount Chambers 1 (MC1 TOP) and (MC1 BASE) Midden 188 7.2.2 Mount Chambers 1 (MCI EXPT) and (MCI EXPB) Midden 194 7.2.3 Mount Chambers 2 (MC2 OVERHANG) and (MC2 TOP) Midden 194 7.2.4 Chambers Gorge 1 (CGI) Midden 197 7.2.5 Chambers Gorge 2 (MD3) Midden 200 7.3 Brachina Gorge Middens 201 7.3.1 Brachina Gorge 1 (BR1 TOP), (BR1 MIDDLE) and (BR1 BASE) Midden 201 7.3.2 Brachina Gorge 7 (BR7 TOP), (BR7 MIDDLE), (BR7 BASE) and Brachina Gorge 2 (BR2) Midden 207 7.3.3 Brachina Gorge 3 (BR3 TOP), (BR3 MIDDLE) and (BR3 BASE) Midden 212 7.3.4 Brachina Gorge 4 (BR4) Midden 215 7.4 Conclusion 218

CHAPTER 8: METHODOLOGICAL ISSUES AND INTERPRETATION OF HOLOCENE VEGETATION 8.1 Introduction 220 8.2 Methodological Issues 220 8.2.1 Accelerator Mass Spectrometry (AMS) Radiocarbon Chronology 220 8.2.2 Comparison Between Midden Pollen and Modern Vegetation 225 8.2.3 Comparison Between Fossil Pollen and Macrofossils in Middens 233 8.3 Regional Patterns in Pollen Data for Individual Taxa 238 8.4 Overview of Holocene Vegetation Communities 247 8.5 Conclusion 250 Vlll

CHAPTER 9: LONG-TERM VEGETATION CHANGE AND HOLOCENE PALAEOCLIMATES FROM STICK-NEST RAT MIDDEN RECORDS 9.1 Long Term Change in Flinders Ranges Vegetation 251 9.2 Causes of Vegetation Change: climate, fire and biogeographical parameters 251 9.3 Holocene Palaeoclimates in the Flinders Ranges 257 9.4 Conclusion 258

CHAPTER 10: CONCLUSIONS AND FURTHER WORK 259

REFERENCE LIST 262

APPENDIX 1: WEIGHTS OF MIDDEN AND SORTED FRACTIONS 283

APPENDIX 2: STANDARD POLLEN EXTRACTION PROCEDURE 284

APPENDIX 3: IDENTIFICATION FEATURES OF POLLEN GRAINS 285

APPENDIX 4: POLLEN COUNTS FOR MIDDENS AND TRAPS 293

APPENDDC 5: POLLEN COUNTS GROUPED INTO VEGETATION CATEGORIES 319

APPENDDC 6: PREPARATION OF POLLEN CONCENTRATES FOR AMS DATING 328

APPENDIX 7: CLUSTER ANALYSIS RESULTS AND DENDROGRAMS 329

APPENDIX 8: CHI-SQUARE STATISTICS FOR POLLEN TRAP DATA 332

APPENDIX 9: MACROFOSSIL:POLLEN INDEX AND SORENSON SIMILARITY INDEX VALUE CALCULATIONS 335

APPENDIX 10: PLANT LIST FOR SPECIES RECORDED AT MIDDEN SITES IN THE NORTHERN AND CENTRAL FLINDERS RANGES 350 ix

LIST OF PLATES

1.1: An example of a stick nest built by Leporillus conditor (Greater-Stick-nest rat) from Monarto South Australia. 2 1.2: Sample of a midden showing the amberat coating that preserves macrofossils and pollen. 2 3.1: Riverine woodland community dominated by Eucalyptus camaldulensis in Brachina Creek at the western end of Brachina Gorge in the central Flinders Ranges. 46 3.2: Low woodland in the Arkaroola-Mount Painter Sanctuary dominated by Callitris columellaris. 46 3.3: Low open woodland dominated by Acacia spp. on lower slopes and Callitris on higher ground near Arkaroola Creek in the Arkaroola-Mount Painter Sanctuary. 48 3.4: A mix of tall shrubland and open shrubland with Acacia and Eremophila spp., on a rocky slope in the Arkaroola-Mount Painter Sanctuary. 48 3.5: Chenopod shrubland community on the plain flanking the western side of the Flinders Ranges outside Brachina Gorge. 49 3.6: Upland tussock grassland community in the Arkaroola-Mount Painter Sanctuary. 49 4.1: Vegetation transect run parallel to the slope contour outside a midden cave site in the Arkaroola-Mount Painter Sanctuary. Species inventories and the foliage cover of species intercepting the transect tape were recorded. 59 4.2: An example of a pollen trap used for both the west-east transect and midden cave pollen studies. 59 4.3: Pollen trap located along the west-east transect across the Flinders Ranges in the Arkaroola-Mount Painter Sanctuary. 60 4.4: Pollen trap anchored inside a midden cave site in the Arkaroola-Mount Painter Sanctuary. 60 5.1: Vegetation on the slopes of Haematite Hill 1 cave site. 115 5.2: Vegetation cover on the rocky slope outside North Well Creek 1 cave site. 115 5.3: Triodia irritans (spinifex) covered slope outside North Well Creek 2 cave site. 120 5.4: Vegetation surrounding the plunge pool outside Waterfall 1 cave site. 120 5.5: Vegetation outside Arkaroola 1 cave site. Acacia victoriae, Eremophila spp. and a ground cover of herbs were common on the rocky slope. 126 5.6: Riverine woodland along Arkaroola Creek downslope from Radium Creek 1 cave site. 126 5.7: Steep sparsely vegetated rocky slope outside Radium Creek 2 cave site. Melaleuca glomerata are present at the base of the slope. 130 X

5.8: Eremophila freelingii and Triodia irritans commonly found on the rocky slope outside Radium Creek 3 midden cave site. 130 5.9: Eremophila freelingii shrubs growing outside the cave site at Oppaminda Creek. 138 5.10: Sparse vegetation cover outside MCI cave site on the south western face of Mount Chambers. 138 5.11: Vegetation (downslope from MCI) outside MC2 cave site on the south western face of Mount Chambers. 143 5.12: Sparse vegetation including Ptilotus obovatus and Eremophila freelingii on the scree slope outside CGI midden site. 143 5.13: Vegetation downslope of MD3 cave site at the eastern end of Mount Chambers Gorge including Eucalyptus camaldulensis, Dodonaea viscosa and ground cover including Ptilotus obovatus and Stipa spp. 151 5.14: Eremophila freelingii, Dodonaea viscosa and Alectryon oleifolium shrubs growing on the slope outside BR1 cave midden site. 151 5.15: Shrubland dominated by Eremophila spp. and ground cover of Ptilotus obovatus and herbs at the western end of Brachina Gorge looking out from the entrance ofBR2/7 cave site. 158 5.16: Steep rocky slope and vegetation cover outside BR3/4 midden cave site. Casuarina cristata was growing outside the cave entrance. 158 LIST OF FIGURES

1.1: Leporillus conditor (Greater Stick-nest Rat) and the former habitat range of Leporillus spp. on the mainland in relation to the extent of the semi-arid and arid zones. 4 3.1: Regional location map of the Flinders Ranges in South Australia. 34 3.2: Location of the summer rainfall/winter rainfall boundary in Australia and proximity to Lake Frome (after Singh 1981: 421). 36 4.1: Location of key stick-nest rat midden sites in the central and northern Flinders Ranges: Arkaroola, Mount Chambers Gorge and Brachina Gorge. 55 5.1: Location of west-east transects for the regional modern pollen trap study at a) Arkaroola b) Mount Chambers Gorge and c) Brachina Gorge. 5.2: Pollen diagram of 1995 and 1995 traps from a west-east transect through the Arkaroola-Mount Painter Sanctuary in the northern Flinders Ranges. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.2. 77 xi

5.3: West-east transect in the Arkaroola-Mount Painter Sanctuary showing pollen trap sites and associated vegetation communities. For sampling methodology see text and for locations of traps see Figure 5.1. 79 5.4: Pollen diagram of 1995 and 1996 traps along a west-east transect through Mount Chambers Gorge in the central Flinders Ranges. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1 % are listed in Table 5.5. 84 5.5: West-east transect through Mount Chambers Gorge showing pollen trap sites and associated vegetation communities. For sampling methodology see text and for locations of traps see Figure 5.1. 86 5.6: Pollen diagram of 1995 and 1996 traps along a west-east transect through Brachina Gorge in the central Flinders Ranges. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.8. 90 5.7: West-east transect through Brachina Gorge showing pollen trap sites and associated vegetation communities. For sampling methodology see text and for locations of traps see Figure 5.1. 93 5.8: Cluster analysis of modern pollen rain from all 1995 trap from west-east transects in the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge. 96 5.9: Cluster analysis of modern pollen rain from all 1996 trap from west-east transects in the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge. 97 5.10: Cluster analysis of all 1995 and 1996 traps from west-east transects in the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge. 99 5.11: Histograms of Australian Bureau of Meteorology monthly rainfall data (1994- 1996) for six regional stations in the central and northern Flinders Ranges. 101 5.12: Pollen diagram of 1995 and 1996 traps (2 and 3) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour on Haematite Hill, Arkaroola-Mount Painter Sanctuary. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. Ill 5.13: Pollen diagram of 1995 and 1996 traps (6 and 7) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at North Well Creek 1, Arkaroola-Mount Painter Sanctuary. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. 114 5.14: Pollen diagram of 1995 and 1996 traps (4 and 5) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at North Well Creek 2, Arkaroola-Mount Painter Sanctuary. Pollen sum xii

equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. 117 5.15: Pollen diagram of 1995 and 1996 traps (9) from outside the midden overhang and foliage cover of species along transects parallel to the slope contour site at Waterfall 1, Arkaroola-Mount Painter Sanctuary. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. 118 5.16: Pollen diagram of 1995 and 1996 traps (19 and 20) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Arkaroola 1, Arkaroola-Mount Painter Sanctuary. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. 121 5.17: Pollen diagram of 1995 and 1996 traps (17 and 18) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Radium Creek 1, Arkaroola-Mount Painter Sanctuary. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. 123 5.18: Pollen diagram of 1995 and 1996 traps (15 and 16) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Radium Creek 2, Arkaroola-Mount Painter Sanctuary. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. 125 5.19: Pollen diagram of 1995 and 1996 traps (12 and 13) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Radium Creek 3, Arkaroola-Mount Painter Sanctuary. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. 128 5.20: Pollen diagram of 1995 and 1996 traps (21 and 22) from inside and outside the midden cave site at and foliage cover of species along transects parallel to the slope contour at Oppaminda Track, Arkaroola-Mount Painter Sanctuary. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. 131 5.21: Pollen diagram of 1995 and 1996 traps (1 and 2) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Mount Chambers 1 on the south western side of Mount Chambers. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.16. 137 5.22: Pollen diagram of 1995 and 1996 traps (3 and 4) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Mount Chambers 2 on the south western side of Mount Chambers. xiii

Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.16. 140 5.23: Pollen diagram of 1995 and 1996 traps (5 and 6) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Chambers Gorge 1 inside the gorge. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.16. 142 5.24: Pollen diagram of 1995 and 1996 traps (7 and 8) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Chambers Gorge 2 (MD3) at the eastern end of the gorge. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.16. 145 5.25: Pollen diagram of 1995 and 1996 traps (1 and 10) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Brachina 1 at the western end of Brachina Gorge. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.18. 150 5.26: Pollen diagram of 1995 and 1996 traps (5 and 6) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Brachina 2/7 at the western end of Brachina Gorge. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.18. 153 5.27: Pollen diagram of 1995 and 1996 traps (7 and 8) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Brachina 3/4 at the western end of Brachina Gorge. Pollen sum equals the total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.18. 155 6.1: Location of stick-nest rat midden sites in the Arkaroola-Mount Painter Sanctuary. Site numbers correspond to those referred to in the midden descriptions in Table 6.1 through to 6.9. 166 6.2: Pollen diagram for middens from the Arkaroola-Mount Painter Sanctuary. Taxa recorded at levels of less than 1% in all middens from this group are listed in Table 6.12. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. 177 6.3: Pollen diagram for Radium Creek 3 midden. Taxa recorded at less than 1% in both top and base samples are listed in Table 6.12. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. 6.4: Pollen diagram for Haematite Hill 1 midden. Taxa recorded at less than 1% in both top and base samples are listed in Table 6.12. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. XIV

6.5: Pollen diagram for Arkaroola 1 midden. Taxa recorded at less than 1% in all sub-samples of the midden (top, middle and base) are listed in Table 6.12. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. 185 7.1: Location of stick-nest rat midden sites at Mount Chambers Gorge. Site numbers correspond to those referred to in the midden descriptions in Table 7.1 through to 7.4. 189 7.2: Pollen diagram for middens at Mount Chambers Gorge 1. Taxa recorded at less than 1% are listed in Table 7.6. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. 195 7.3: Pollen diagram for middens Mount Chambers Gorge 2. Taxa recorded at less than 1% are listed in Table 7.6. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. 198 7.4: Pollen diagram for middens inside Chambers Gorge. Taxa recorded at less than 1% are listed in Table 7.6. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. 198 7.5: Location of stick-nest rat middens from Brachina Gorge. Site numbers correspond to those referred to in the midden descriptions in Table 7.8 through to 7.11. 202 7.6: Pollen diagram of Brachina Gorge 1 midden. Taxa recorded at less than 1% are listed in Table 7.13. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. 208 7.7: Pollen diagram of Brachina Gorge 2 and Brachina Gorge 7 middens. Taxa recorded at less than 1% are listed in Table 7.13. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. 7.8: Pollen diagram of Brachina Gorge 3 midden. Taxa recorded at less than 1% are listed in Table 7.13. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. 214 7.9: Pollen diagram of Brachina Gorge 4 midden. Taxa recorded at less than 1% are listed in Table 7.13. Pollen sum equals total pollen counted. A dot indicates presence of macrofossils of the pollen taxon in the midden. 216 8.1: Scatter plots of AMS dates on faecal pellets and leaf macrofossils from top and base samples of Brachina Gorge 1 and Brachina Gorge 3 middens. 222 8.2: Brachina Gorge 4 midden displays apparent layering. AMS dates on faecal pellets from top and base samples and AMS dates on leaf macrofossils and pollen concentrates at 10cm intervals are illustrated on the schematic diagram and scatter plot. 223 XV

8.3: Percentages of key pollen taxon recorded in the northern and central Flinders Ranges from stick-nest rat middens located in the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge. 240 8.4: Composition of Holocene vegetation communities from fossil pollen records from stick-nest rat middens at study sites in the northern and central Flinders Ranges. 248 8.5: Summary of modern pollen from traps along west-east transects through the Flinders Ranges at Arkaroola, Mount Chambers Gorge and Brachina Gorge. LIST OF TABLES

5.1: Location and recovery rate of pollen traps from the 1995 and 1996 west-east transects from the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge. Locations of traps are shown on Figure 5.1 (a), (b) and (c) respectively. 75 5.2: Pollen taxa recorded at levels of less than 1% in traps from the west-east transect modern pollen study in the Arkaroola-Mount Painter Sanctuary. 78 5.3: Comparison of taxa recorded in the modern pollen and vegetation at trap sites along the west-east transect at Arkaroola. Data was used to calculate a Sorenson Similarity Index for each site. 80 5.4: Sorenson Similarity Index for comparison of taxa represented in the modern pollen rain and vegetation cover at trap sites along the Arkaroola-Mount Painter Sanctuary west-east transect for the sampling time of vegetation. 82 5.5: Pollen taxa recorded at levels of less than 1% in traps along the west-east transect modern pollen study at Mount Chambers Gorge. 85 5.6: Comparison of taxa recorded in the modern pollen rain and vegetation at trap sites along the west-east transect at Mount Chambers Gorge. Data was used to calculate a Sorenson Similarity Index for each site. 87 5.7: Sorenson Similarity Index for comparison of taxa represented in the modern pollen rain and vegetation cover at trap sites along the west-east transect at Mount Chambers Gorge. 88 5.8: Pollen taxa recorded at levels of less than 1% in traps from the west-east transect modern pollen study at Brachina Gorge. 91 5.9: Comparison of taxa recorded in the modern pollen and vegetation at trap sites along the west-east transect at Brachina Gorge. Data was used to calculate a Sorenson Similarity Index for each site. 94 5.10: Sorenson Similarity Index for comparison of taxa represented in modern pollen rain and vegetation cover at trap sites along the west-east transect at Brachina Gorge. 95 XVI

5.11: Location and name of Australian Bureau of Meteorology regional rainfall stations in the northern and central Flinders Ranges. 100 5.12: Representation of pollen taxon in the modern pollen rain from the northern and central Flinders Ranges. 103 5.13: Location and recovery rate of pollen traps from the 1995 and 1996 study from midden sites in the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge. 109 5.14: Taxa recorded at levels of less than 1% in cave traps from midden sites in the Arkaroola-Mount Painter Sanctuary for 1995 and 1996 sampling periods. 132 5.15: Comparison of taxa recorded in the modern pollen and vegetation at midden cave sites in the Arkaroola-Mount Painter Sanctuary. Data was used to calculate a Sorenson Similarity Index for each site. 134 5.16: Taxa recorded at levels of less than 1% in cave traps from midden sites on Mount Chambers and in Mount Chambers Gorge for 1995 and 1996 sampling periods. 146 5.17: Comparison of taxa recorded in the modern pollen and vegetation at midden cave sites at Mount Chambers Gorge. Data was used to calculate a Sorenson Similarity Index for each site. 148 5.18: Taxa recorded at levels of less than 1% in cave traps from midden sites in Brachina Gorge for 1995 and 1996 sampling periods. 150 5.19: Comparison of taxa recorded in the modern pollen and vegetation at midden cave sites in Brachina Gorge. Data was used to calculate a Sorenson Similarity Index for each site. 157 5.20: SI values for a comparison between vegetation cover and modern pollen rain at cave traps from stick-nest rat midden sites in the Arkaroola-Mount Painter Sanctuary. 159 5.21: SI values for a comparison between vegetation cover and modern pollen rain at cave traps from stick-nest rat midden sites on Mount Chambers and in Mount Chambers Gorge. 159 5.22: SI values for a comparison between vegetation cover and modern pollen rain at cave traps from stick-nest rat midden sites at Brachina Gorge. 159 6.1: Description of Haematite Hill 1 midden and environmental features at the cave site. 167 6.2: Description of North Well Creek 1 midden and environmental features at the cave site. 168 6.3: Description of North Well Creek 2 midden and environmental features at the cave site. 169 6.4: Description of Waterfall 1 midden and environmental features at the cave site. 170 xvii

6.5: Description of Radium Creek 1 midden and environmental features at the cave site. IV1 6.6: Description of Radium Creek 2 midden and environmental features at the cave site. 172 6.7: Description of Radium Creek 3 midden and environmental features at the cave site. 173 6.8: Description of Arkaroola 1 midden and environmental features at the cave site. 174 6.9: Description of Oppaminda Track 1 midden and environmental features at the cave site. 175 6.10: AMS radiocarbon dates for middens from the Arkaroola-Mount Painter Sanctuary. 176 6.11: Macrofossil assemblages for middens from the Arkaroola-Mount Painter Sanctuary. 6.12: Pollen taxa recorded at levels of less than 1% in the pollen sum for middens from the Arkaroola-Mount Painter Sanctuary. 186 7.1: Description of Mount Chambers 1 midden and environmental features at the cave site. 190 7.2: Description of Mount Chambers 2 midden and environmental features at the cave site. 191 7.3: Description of Chambers Gorge 1 midden and environmental features at the cave site. 192 7.4: Description of Medlin Site 3 midden and environmental features at the cave site. 193 7.7: Macrofossil assemblages for middens from Mount Chambers Gorge. 196 7.6: Pollen taxa recorded at levels of less than 1% in the pollen sum for middens from Mount Chambers Gorge. 199 7.5: AMS radiocarbon dates for middens from Mount Chambers Gorge. 200 7.8: Description of Brachina 1 midden and environmental features at the cave site. 203 7.9: Description of Brachina 2 and Brachina 7 middens and environmental features at the cave site. 204 7.10: Description of Brachina 3 midden and environmental features at the cave site. 205 7.11: Description of Brachina 4 midden and environmental features at the cave site. 206 7.12: AMS radiocarbon dates for middens from Brachina Gorge. 210 7.13: Pollen taxa recorded at levels of less than 1% in the pollen sum for middens from Brachina Gorge. 217 7.14: Macrofossil assemblages for middens from Brachina Gorge. 8.1: Comparison of taxa recorded in midden pollen and vegetation at midden cave sites at Arkaroola. 227 xviii

8.2: Comparison of taxa recorded in midden pollen and vegetation at midden cave sites at Mount Chambers Gorge. 230 8.3: Comparison of taxa recorded in midden pollen and vegetation at midden cave sites at Brachina Gorge. 231 8.4: Macrofossil'.Pollen Indices for middens from the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge. 234 8.5: Macrofossil:Pollen Index for Early, Middle and Late Holocene middens. 236 8.6: Summary of representation of key taxa in the modern pollen rain, midden pollen and macrofossils. 238 -1-

Chapter 1: Introduction: Thesis and Aims

1.1 Palaeoecology of the Semi-Arid Zone: the issues, questions and problems

Palaeoecological evidence is widely recognised as being crucial to our understanding of long term biogeographic and climatic processes. This knowledge in Australia is biased towards well-watered parts of the continent. For example, pollen preserved in lake and swamp sediments has traditionally provided the main source of evidence for reconstructing vegetation histories over time. Approximately 70% of the continent is semi-arid and arid and these dry environments are not conducive to the preservation of this fossil material, except in some salt lakes which have records very regional in character. While the increasing body of geomorphic evidence from inland areas is providing palaeoclimatic data, it is by its nature broad scale. Lack of Quaternary pollen and macrofossil records from inland areas indicates that the precise nature of climate change and biotic responses remain unknown for a large part of the continent. Elucidating the palaeoecology and palaeoclimates for this significant area in the Southern Hemisphere will provide an important source of data on global climate change.

1.2 The Evidence

Native stick-nest rat (Leporillus conditor and Leporillus apicalis) middens offer a unique source of biogeographical evidence for arid and semi-arid environments. Stick- nest rats assembled mounds of sticks, leaves, grass and stones that were collected within a 100 metre foraging range of the nest site, and subsequently compacted together to form nests (Plate 1.1) (Copley 1988; Tunbridge 1991). There were nesting chambers constructed inside the nest that were lined with grass, soft leaves and chewed bark strips and linked to the outside via a number of tunnels (Copley 1998; D. Kennett pers.comm. 1992). Once assembled, sections of the nest became cemented by a covering of the rat's viscous urine. This coating hardened into a black tarry substance known as amberat, sealing and preserving the faecal pellets, plant fragments, vertebrate remains, stones, dust and pollen that accumulated in the nests over time. Nests in this condition are referred to as middens (Plate 1.2). The appearance of nests and middens range from dome shaped nests in overhangs or caves to token gatherings of twigs and stones on flat open terrain (Watts and Eve 1976; Watts and Aslin 1981; Copley 1988; Tunbridge 1991) and the size appears dependent upon available nesting material at the time of construction, location on the landscape, age and length of occupation (Tunbridge 1991).

When the rats became extinct on the mainland in the years following European settlement, they left a legacy which provides the opportunity to examine vegetation -2-

Plate 1.1: An example of a stick nest built by Leporillus conditor (Greater-Stick-nest rat) from Monarto South Australia.

Plate 1.2: Sample of a midden showing the amberat coating that preserves macrofossils and pollen. -3-

change over a range of time scales, from Pleistocene to post-European. The former distribution of Leporillus (Figure 1.1) in the southern part of the semi-arid and arid zone overlaps extensive areas for which there is virtually no Late Quaternary palaeoecological evidence.

1.3 The Thesis: Methodological and Taphonomic Issues

Previous work with stick-nest rat middens has demonstrated the great potential of this material for palaeoecological research (Green et al. 1983; Nelson et al. 1990; Pearson and Dodson 1993; McCarthy 1993; McCarthy et al. 1996; Allen 1996; Head et al. 1997; Pearson 1998) for example, in initial work I used midden records to reconstruct a generalised Holocene record for the Flinders Ranges. Singh and Luly's (1991) reliance on the summer/winter rainfall distribution to explain vegetation response throughout the Holocene in the semi-arid zone was not entirely supported by the first study of stick- nest rat middens (McCarthy et al. 1996). It became apparent there was a need for a regional project that systematically covers a significant geographic area while addressing unresolved methodological questions. These questions relate to the spatial and temporal resolution of middens with reference to accumulation rates and chronologies of deposits, the possibility of reworking of material, and identification of source and recruitment processes of pollen into the midden matrix. These factors have to be resolved in order to increase the integrity of palaeoecological and palaeoclimatic interpretations based on these records. The emphasis on understanding the processes of midden construction is important in the Australian context as fossil pollen is more successfully preserved in middens while the macrofossil record is poor compared to North American Neotoma middens. Interpretation of palaeovegetation communities is also more difficult as there is high spatial variability in arid zone vegetation communities and a lack of distinct altitudinal zonation. This contrasts with North America where analysis of Neotoma spp. middens has yielded very precise information about the dynamics of desert plant communities from 40 000 years to the present, providing insights about mountain top biota and island biogeographic theory (see examples in Betancourt et al. 1990b).

1.4 Thesis Objectives

This project uses Leporillus middens to provide a systematic regional study of Holocene vegetation histories and palaeoclimates in the Central and Northern Flinders Ranges. The temporal resolution of midden records is coarse, compared to traditional pollen core data, however this factor has been offset by the extensive spatial coverage provided by middens. Key study sites are located in the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge in South Australia. -4-

Arid and semi-arid zone.

Former distribution of stick-nest rats. t

«)• uf 120- t«r [

Figure 1.1: Leporillus conditor (Greater Stick-nest Rat) and the former habitat range of Leporillus spp. on the mainland in relation to the extent of the semi-arid and arid zones. -5-

There are four broad aims which are addressed via the following specific objectives: 1) Increase knowledge of modern pollen deposition in semi-arid environments and likely sources for midden pollen: a) Investigate sources of pollen recruited into middens and at study sites, by conducting regional and local modern pollen rain studies and comparing this data with present day vegetation communities. 2) Understand taphonomy of stick-nest rat midden deposits: a) Measure the time span over which large midden deposits accumulate to provide well dated palaeoecological records from single sites. b) Refine the temporal resolution and palaeoecological interpretations of middens using Accelerator Mass Spectrometry Radiocarbon dating of individual components ie. macrofossils, faecal pellets and pollen concentrates from deposits. c) Assess the palaeoecological significance and potential of pollen and macrofossil assemblages after consideration of midden sample size and variability within middens and from middens of a similar age. 3) Elucidate a regional Holocene vegetation history for the Flinders Ranges: a) Assess the potential for detecting spatial and/or temporal variation in midden records from the regional study. b) Use the spatial and temporal coverage provided by the middens to reconstruct Holocene vegetation change for the northern and central Flinders Ranges. 4) Reconstruct Holocene palaeoclimates from stick-nest rat midden palaeoecological records: a) Compare palaeoclimatic signals inferred from midden records to other lines of evidence including traditional pollen cores, lake levels and geomorphic evidence. b) Test movements of significant climatic parameters such as the summer/winter rainfall in the semi-arid zone using stick-nest rat midden records.

1.5 Chapter Outline

Chapter 2 outlines the research context for this project. A review of the field of midden analysis from other parts of the world is presented that addresses specific taphonomic and methodological issues and the role of pollen in midden analysis. The focus is on pollen, as the macrofossil content of Australian middens is minimal and not well preserved. Holocene palaeoenvironments of Australia are reviewed, concluding with a summary of the context in which this project sits and the important contribution that palaeoeoclogical evidence from stick-nest rat middens makes to further our understanding of the complex interactions between climate and biogeographic changes in the arid and semi-arid zones. -6-

An overview of the regional setting of the Flinders Ranges is presented in Chapter 3. There is a general discussion of the semi-arid environment in terms of factors that influence the composition and structure of vegetation (topography, geology, aspect, rainfall, European impacts). This provides the context for a discussion of vegetation communities and environmental features of midden study sites.

Chapter 4 describes the methodological design of the project. The methodologies are discussed within the context of my research objectives and include: a field work component (midden surveys, midden excavation, vegetation surveys); modern pollen rain study (trap locations, vegetation quadrat data); laboratory work (midden sub- sampling, processing for pollen and macrofossils, pollen counting and analysis); radiocarbon dating of macrofossils and pollen concentrates; and statistical analysis of data sets.

Chapter 5 presents the results of regional and local modern pollen studies from traps located along west-east transects across the Flinders Ranges and at midden cave sites. This is followed by a discussion of this data in the context of temporal and spatial variability in the modern pollen rain and the important implications this holds for the interpretation of fossil pollen recovered from middens.

Chapter 6 and Chapter 7 present the fossil pollen and macrofossil assemblages from midden sites from the northern ranges and central ranges respectively.

Chapter 8 begins with a discussion of taphonomy and chronologies of midden deposits. Overall findings are presented on the basis of Accelerator Mass Spectrometry radiocarbon dates on leaf macrofossils, faecal pellets and pollen. Relationships between midden pollen and macrofossils, and recruitment of pollen into middens, is explored via macrofossil:pollen index values and comparing present day vegetation and modern pollen rain with midden pollen assemblages. The end result being to increase our understanding of the integrity of each fossil record in terms of usefulness for palaeoecological and palaeoclimatic reconstructions. A Holocene vegetation history derived from the pollen and plant macrofossil assemblages is then presented. To tease out the complexity of the Holocene story there is a comparison of northern versus southern sites of similar ages, and a comparison of site records spanning the Holocene. Both of these contribute to a discussion of Holocene vegetation within the regional context of the Flinders Ranges and inferred palaeoclimates from different sites in Chapter 9. -7-

Chapter 2: The Research Context

2.1 Introduction

Three main sections in this chapter address the field of midden analysis, gaps in our understanding of Holocene palaeoenvironments in Australia, and the contribution that evidence from Leporillus spp. middens can make to increase our knowledge of Holocene palaeoenvironments. The chapter begins with a literature review that addresses two main themes in midden analysis; methodological and taphonomic issues and the palaeoecological information compiled from these records. As Anderson and Van Devender (1991) stated, it is important to resolve these issues to increase the integrity and reliability of interpretations of fossil material from animal middens. I am placing importance on methods and taphonomy in midden research as there has been less attention to this area because of the abundant macrofossil record in Pleistocene and Holocene Neotoma middens. The second section reviews Holocene palaeoenvironments of Australia based on different lines of evidence; lake levels, pollen records and dune and fluvial records. The final section presents a summary of the context in which this project sits and the gaps in the knowledge of semi-arid palaeoecology which it can fill.

2.2 Midden Analysis

A general introduction to Neotoma (packrats) and the field of midden analysis is presented. This is followed by examples of Neotoma midden studies that have contributed to a comprehensive palaeoecological history of desert areas in North America. A review of midden research in Africa and the Middle East highlights the different approaches and traditions in midden analysis from work in North America. African and Middle Eastern midden studies have emphasised methodological issues. Australian work is then described, setting the context for this project.

There are some similarities and differences between Leporillus spp. midden studies from the semi-arid and arid zones of Australia and Neotoma spp. studies. Leporillus studies have been directed towards understanding taphonomy of middens, methodological issues and the palaeoecological significance of multiple sources of evidence (ie. midden pollen, macrofossils and insect assemblages) from the beginning. Plant macrofossils are less abundant and less adequately preserved in stick-nest middens and collecting behaviour of the rat is less known compared to Neotoma middens. Procaviidae (hyrax) are herbivores that are not known to carry food or other plant remains into their shelter but browse within 50m-100m (Scott 1990) but in Hyrax middens from the Middle East, plant fragments are incorporated into middens (Fall et -8-

al. 1990). Faecal pellets accumulate in hyrax refuges and form extensive middens when cemented by the hyrax's concentrated urine in contrast to Petromuridae (dassie rat) middens that contain plant remains and faecal pellets, similar to Neotoma middens. Dassie rats are known to take plant material back to shelters in order to feed (Scott 1990).

2.2.1 Neotoma Midden Analysis - the beginnings

Packrat middens are waste piles of organic material, mixed with faecal pellets and cemented with urine (Spaulding 1992). Each deposit may contain preserved seeds, twigs, leaves and fruits. This is material collected around the immediate vicinity of the den, and hence regarded as precise vegetation data (Spaulding 1992). A urine coating (amberat) preserves the midden contents (Wells 1976; Cole 1990; Spaulding et al. 1990; Betancourt et al. 1990b) and also has an adhesive property enabling the midden to be glued to a surface inside dry rock shelters or crevices (Spaulding et al. 1990).

Horton and Wright (1944) described a packrat nest as a shelter of sticks built around a supporting bush. Wells (1976) suggested that the stick nest and midden are separate entities. The outer stick structure continues to accumulate while the nest is occupied. There is an indurated finer plant debris layer underneath consisting of a network of chambers and passages, referred to as the interior middens. Occasionally these structures will be cleaned out by pushing refuse through entrances, forming large exterior middens (Wells 1976; VanDevender 1987; King and VanDevender 1977). In caves and overhangs, the architecture of stick nests degenerates to middens of plant fragments and faecal pellets (Wells 1976).

Wells (1976) addressed the ecology and behaviour of packrats in a pioneering study. He suggested that the rat forages for building materials and a food supply within a 100 metre radius of the nest. Cole (1990) suggested that midden contents reflected plant species within 10-30 metres of the nest site, Van Devender and Wiens (1993) suggested a 30 metre radius, and Spaulding (1990) 30-50 metres. This material is incorporated into the nest, contributing to part of the macrofossil record (Wells 1976; King and VanDevender 1977) as the rats collect more food than is consumed (Wells 1976; King and VanDevender 1977; Dial and Czaplewski 1990).

Announcement of the scientific potential of Neotoma middens in the semi-arid and arid zones of America occurred in the 1960s (Wells and Jorgenson 1964; Webb and Betancourt 1990; Nowak et al. 1994). Macrofossil assemblages and pollen have provided information for the detailed reconstruction and distribution of Late Quaternary plant communities and individual species from mountain, upland and basin topography -9-

in deserts of Western North America and northern Mexico, specifically the Chihuahuan, Sonoran, Mojave, Great Basin and The Colorado Plateau Desert regions (Betancourt and Davis 1984; Elias 1990; Baynes 1991; O'Rourke 1991; Van Devender etal. 1991). Emphasis has been placed on desert scrub, pygmy conifer woodlands, sub-alpine and montane woodlands, Ponderosa pine and grassland communities (Betancourt et al. 1990a). Records from midden analysis have also been used for palaeoclimatic interpretations, evidence of displacements of vegetation types along altitudinal and latitudinal gradients and mammal faunal changes that have occurred in desert environments.

2.2.2 Contribution of Neotoma Midden Studies to the Palaeoecology of Desert Areas in the USA

Within this foundation work, there has not been a great deal of research into methodological issues questions on midden analysis. Neotoma middens as old as 40 000 years to the present have provided long term records of vegetation histories for extensive areas throughout North American Deserts. The macrofossil assemblages represent local vegetation at discrete time periods and regional sequences can be constructed from different middens of different ages (Thompson 1988). Given these conditions, US research has focussed on documenting long term palaeoenvironmental histories. It was not until the late 1970s that direction shifted towards inclusion of midden pollen analysis and attention to methodological complexities in the nature of fossil middens such as biases reflected in the composition of macrofossil and pollen records, age and structure of the midden, relationship between species abundance in the midden compared to the plant communities from which midden contents were derived.

Major contributions to the palaeoecology of the south west USA by Neotoma midden studies are demonstrated in this section with some examples of this research. It is agreed upon in midden literature that Late Pleistocene desert woodlands and chaparral have been replaced by desert scrub throughout large areas of North American deserts, but debate remains over the extent of treeless communities during the Last Glacial (Betancourt et al. 1990a). The Late Pleistocene was dominated by pinyon juniper and oak trees (Pygmy Conifer Woodlands) that are now found at desert elevations. These species were stable during the Mid-Late Pleistocene until 11 000 years B.P., when the pinyon pines vacated the lowlands but remained at desert elevations into the Early Holocene. During the Late Pleistocene, Sub-Alpine and Montane Forests dominated areas now occupied by Pygmy Conifer Woodlands. The Pleistocene history of Ponderosa Pine Woodlands has been re-interpreted after the identification of different pine species in macrofossil assemblages (i.e. Limber versus Ponderosa pine) (Betancourt et al. 1990a). Information on grassland communities (proportion of C^ to -10-

C+ grasses) has been useful for palaeotemperature indicators. Midden records have shown that the proportion of C^ grasses increases with elevation and a north trending gradient (Betancourt et al. 1990a).

Nowak et al. (1994) presented a 30 000 year record of vegetation dynamics in the Great Basin Southwest US, based on plant macrofossils from Neotoma middens. The context of this research, based within a diverse topographic setting (basin to mountain range topography), was an excellent opportunity to study plant responses to climatic variations. It was acknowledged that an advantage of using middens provided an investigation into vegetation dynamics at a local scale (Nowak et al. 1994).

Neotoma middens from Ragged Top, South-Central Arizona provided a record from approximately 14 000-5 000 years B.P. to investigate the developmental history of the flora and information on the timing of isolation of relicts. The macrofossil records showed that the Ragged Top flora shifted composition in the Holocene as woodland species died out and Sonoran desert scrub species arrived at different times (Van Devender and Wiens 1993).

Comparison of pollen versus macrofossil records has also been a focus in Neotoma research. A detailed 2 000 year pollen record from Lower Pahranagat Lake, Southern Nevada demonstrated the xeric nature of climate during the Late Holocene and the record could detect a shift in summer rainfall since 1 500 years BP. that encouraged an expansion of grass and pinyon species (Hemphill and Wigand 1994). Macrofossil records have been used to detect the appearance and disappearance of indicator species such as Juniper us, Artemisia and Poaceae, thus an indication of changing plant communities, whereas the pollen records allow reconstruction of long term continuous shifts in plant communities.

Anderson and Van Devender (1995) were the first to study the pollen from Neotoma middens from Sierra-Bacha, north-western Sonora, in northern Mexico. A major conclusion from this work was that pollen analysis is becoming more important in packrat research/midden analysis primarily because it complements macrofossil assemblages. Middens also provide an excellent opportunity for palaeoecological reconstructions at low elevations in arid regions of North America and Mexico (Anderson and Van Devender 1995). The taxonomic resolution of pollen is less than macrofossil identifications however in this study, pollen added some local species and regional types to the fossil records that were not apparent in the macrofossil inventories. Both pollen and macrofossils suggested a wetter and/or cooler Early Holocene (compared to today), mesic Middle Holocene and drying/warming trend in the Late Holocene (Anderson and Van Devender 1995). -11-

Midden analysis has been used within an archaeological context, with references to the occurrence of corn pollen in middens and the implications for cultural settlement in the southwest United States (Betancourt and Davis 1984). Work completed in the Chuska Mountains on the Arizona-New Mexico border showed that pollen and macrofossil analysis from middens did not support interpretations from pollen lake data. It was demonstrated that midden analysis can test the pollen based inference for low-elevation vegetation zones in this region (Betancourt and Davis 1984).

2.2.3 Midden Studies in South Africa and the Middle East

Resolving methodological issues has been the focus in midden studies from South Africa and the Middle East. Likely sources of pollen and macrofossils and the implications of this knowledge for palaeoecological interpretations have been dominant themes. Unlike Neotoma middens, these middens are not always rich in macrofossil floras. Vegetation communities are also different, with less distinct altitudinal zonation of plant species than is characteristic of North American environments. This section outlines the methodological focus in midden research from South Africa and the Middle East by referring to specific examples.

Procavia spp. (Hyrax) and Petromus spp. (Dassie Rat) middens have been analysed in work in South Africa and the Middle East. Petromus middens are restricted to a smaller geographical area compared to hyrax middens (Scott and Cooremans 1992). Scott and Cooremans (1992) investigated the pollen spectra in twenty Hyrax middens, four Dassie Rat middens and one sample of bird guano from the Great Karoo and grassland regions of Orange Free State and Namaqualand on the north west Cape in South Africa. They addressed the differences in the midden pollen spectra from modern Hyrax dung samples (representing the last 1 000 years) to determine whether the pollen reflected local vegetation. Hyrax middens provided a pollen record that was unbiased by diet and showed no clear difference between dung and surface pollen.

Late Holocene hyrax middens (1 570 to 1 120 years BP.) from the Middle East provided a palaeovegetation record and addressed the role of human disturbance in the environment (Fall et al 1990). Main study sites were in the Southern Jordan Valley and the Wadi Araba. Animals randomly sampled vegetation near their dens, and plants and pollen from the middens characterised typical vegetation near the dens at the time of midden construction. Pollen spectra from subsamples of each midden were compared by statistical methods and significant differences within the middens were identified. Similar frequencies of some pollen taxa and different frequencies of others implied different modes of pollen deposition, therefore reflecting the vegetation present at varying distances from the midden. Background pollen rain reflected the extra-local and

* 0009 03254446 7 -12-

regional vegetation and was less biased by foraging of the animal. Fall et al. (1990) concluded plant fragments in the middens were an indication of local vegetation and pollen assemblages were a mixture of regional and local sources.

Analysis of Neotoma cinerea (Bushy Tailed Woodrat) middens from the arid interior of British Columbia Canada, provides a method of completing environmental reconstructions for parts of the landscape not accessible by traditional lake core studies (Hebda et al. 1990). The workers found inconsistencies between the abundance of some species in the vegetation and the amount of pollen in midden samples. Dry habitat taxa were better represented in the middens than in the lake core records for similar time periods (Hebda et al. 1990), suggesting a bias in midden records.

Pollen analysis of Late Holocene middens (most likely Phyllotis xanthopygus, Leaf- eared Mouse) from Argentina, is an expansion of midden research into South America. It was found that midden pollen had a large local component in contrast to regional lake and bog deposits. The sources of pollen included local and regional pollen tracked in on fur, and pollen in faecal pellets (Markgraf et al. 1997).

There are new directions taking place in Neotoma midden research for investigating palaeoclimate reconstructions. Work by Van de Water et al. (1994) measured the stomatal densities and 8^C in Limber pine macrofossils from middens at varying elevations in different states from the South West U.S.A. Shifts in plant physiology and leaf morphology were revealed. Variations in 8^C allowed investigation of the redistribution of carbon between the atmosphere and biosphere during the Last Glacial- Interglacial cycle (Van de Water et al. 1994). Work by Smith et al. (1995) addressed the micro evolutionary changes of Neotoma cinera estimated from the measurement of faecal pellets. Size changes closely tracked regional temperature fluctuations simulated from isotope ratios of plant cellulose from middens. It was found that body size decreased during periods of climatic warming. The study suggests that analysis of faecal pellets from many locations can permit precise quantification of responses to climatic change from the past and into the future (Smith et al. 1995).

Urine from Neotoma middens has been used to increase knowledge of the production history of cosmogenic nuclides, required for geological and archaeological dating. Measures of chlorine-36/chlorine (36Q/C1) ratios in fossil (38 000 years B.P. to present) Neotoma middens from Nevada showed that 36Q/C1 ratios were higher before approximately 11 000 years B.P. This suggests that cosmogenic production rates before the close of the Pleistocene were up to 50% higher than what has been proposed by ^C calibration data (Plummer et al. 1997). Chloride is also a conservative tracer in water and a change in 36Q/C1 ratio could be useful for hydrologic studies. -13-

Analysis of stable isotopes (Deuterium (8D)) in plant cellulose from midden macrofossils has been found to be useful for discriminating climatic change (Long et al. 1990). Measured 5D values from Neotoma middens from the White Mountains, California-Nevada suggested lower temperatures and increased precipitation into the Eary Holocene, relative to the present (Jennings and Elliot-Fisk 1993).

2.2.4 Methodological and Taphonomic Issues in Midden Analysis

In this section I review different factors that need to be considered and assessed, when resolving methodological and taphonomic issues in midden analysis. I highlight historical changes in approach and different traditions of research in different places.

Midden Depositional Histories The depositional history of middens is debated in the literature as considerable disagreement exists over what proportion of packrat dens become indurated and the timing of accumulation of a midden deposit (Betancourt et al. 1990a). Questions over whether Neotoma middens accumulated over hundreds of years (King and VanDevender 1977) or only a few years (Spaulding et al 1983; Jacobson 1988; Spaulding et al. 1990; Thompson 1985) have appeared in earlier work. According to Wells (1976), the time of midden formation is dependent on the size and type of shelter. Packrats gathered a lot of material in short periods of time and, given suitable conditions in a protected site, urine cemented materials into an indurated mass within a few months to a few years with weathering rinds separating different building episodes (Thompson 1985). Other work has suggested that middens can be stratigraphically discontinuous where different samples have to be dated, but unlike traditional palynology from sediment cores, time periods between each sample cannot be inferred. Middens can also be stratigraphically complex if there have been breaks in accumulation or collapse in the structure (Elias 1990; Spaulding et al. 1990; Webb and Betancourt 1990), or where packrats build in a crevice that once contained another midden (Jacobson 1988). These issues can be investigated via rigorous radiocarbon dating of single middens and/or different midden components (i.e. leaves, faecal pellets and pollen).

Interpretation of Midden Pollen Methodological issues which must be addressed include improving the reliability of pollen analysis by investigating pollen sources and recruitment processes into middens; the size of samples used for pollen analysis; consideration of an appropriate pollen sum and an understanding of intra-midden variability. Regional versus local pollen signals need to be addressed as well as the noise in pollen data, the possibility of a seasonal bias in both macrofossil and pollen assemblages, the demonstrated variability of pollen -14-

signals among middens of similar age and site and the representativeness of individual pollen taxa. Each of these issues is addressed in the following section.

Validity and Reliability Davis and Anderson (1986) argued that the reliability of pollen analysis of middens can be evaluated via two kinds of replicate samples: 1) duplicate samples of the same midden unit; 2) different middens of the same age (Davis and Anderson 1986, 1987).

Pollen analysis is frequently related to modern analogues such as plant abundances or climatic variables, but the validity (degree of relationship) is difficult to establish (Birks and Gordon 1985). Correlation of pollen with modern vegetation is a recognised problem in traditional pollen analysis, however middens are different and in some respects problematic. Midden pollen cannot be interpreted if the relationship of pollen to vegetation is not consistent. There is not necessarily a one to one correlation between the proportional occurrence of macrofossil and pollen taxa in the nests/middens and abundance of these taxa in the surrounding vegetation (Wigand and Nowak 1992). The role of the rat, with respect to foraging, midden building and dietary bias are additional factors that influence midden pollen assemblages. Therefore, reliability is important for packrat midden analysis because pollen content is so variable (Davis and Anderson 1986).

Uncertainty over Pollen Sources and Recruitment Knowledge of pollen recruitment to midden deposits is fundamental to achieving rigorous palaeoecological reconstructions. Little is known about the capture of pollen by different plants or plant parts and the consequent effect on the input of fossil pollen into middens. Pollen productivity, dispersal and seasonality of the plant collections incorporated into middens are not constant factors. When interpreting fossil pollen records, it is advantageous to understand the pollen productivity and mode of deposition for species in the vegetation communities (O'Rourke 1991). Filtration of pollen can occur by leaf surface processes such as resinous, glabrous recumbent hairs and/or stellate hairs on plants. There is also the size of the pollen whereby large grains are more effectively filtered by the vegetation than small grains. These characteristics can promote or reduce pollen capture (O'Rourke 1991).

Thompson (1985) and O'Rourke (1991) suggested that pollen recruitment into middens occurs via the following pathways: 1) rat foraging and nesting activities; 2) passive accumulation of windborne material that is dependent upon how open the site is, the length of midden formation and nature of the surrounding vegetation and 3) rat dietary preferences. Davis and Anderson (1986, 1987) suggested additional routes that included: 4) packrat faeces and 5) adherence to packrat pelts, but agreed that airborne -15-

transport and pollen on macrofossils are the most important pathways. It is acknowledged that variations in the relative importance of the other three mechanisms adds to the variability of midden pollen. Finley (1990) stated that it is the interaction between the rat and the environment that is the overall determinant of what and how organic components of middens are transported and how the midden is formed. Pollen introduced by rats and their faecal pellets is subject to bias caused by local collecting and foraging habits and there is argument over the contribution of pollen from this source (King and Van Devender 1977; O'Rourke and Mead 1985). In most instances, pollen in faecal pellets is similar to midden compositions but zoophilous taxa can be elevated. Some studies have indicated that crashed faecal pellets contain a biased coverage of the local plant species in the rats' diet (King and VanDevender 1977; Dial and Czaplewski 1990; Pearson and Dodson 1993). However according to Davis and Anderson (1986; 1987), this source is not too important as most pellets appear to remain intact and deliberate crashing and stirring during processing is required to release the pollen into the midden matrix. Scott and Cooremans (1992) also concluded that Hyrax midden pollen was unbiased by the animal's diet.

Physical parameters of midden deposits may also affect recruitment of pollen taxa. Davis and Anderson (1987) suggested that the mode of pollen transport varies depending on the size and location of the midden. That is, if the midden is exposed (at the front of a cave) there would be greater airborne pollen in contrast to middens located in smaller crevices (Davis and Anderson 1986).

Pollen Counts and Intra and Inter-Midden Variability There should be a large pollen sum to ensure the inclusion of regionally important taxa and entomophilous species. And as locally important species may be low level pollen producers, analysis should be based on high pollen counts and include additional screening for rare pollen types (King and Van Devender 1977).

There is conflict regarding observations of variability within middens and reflection of this in the pollen record. Thompson (1985) suggested that pollen spectra preserved in the middens exhibit little variability within a deposit. This is contrary to Davis and Anderson (1986; 1987) where pollen within the one midden can be variable.

Regional and Local Pollen Signals Midden pollen represents both local and regional vegetation (Anderson and Van Devender 1991). However whether the input of pollen is predominantly local, extra local or regional is in dispute (King and Van Devender 1977; Thompson 1985; Davis and Anderson 1986,1987; Van Devender 1988). Observations have been based on the comparison of pollen taxa recorded in middens with surrounding vegetation and the -16-

local records from well preserved macrofossil assemblages. Attention to the role of Neotoma collection and dietary bias has also been prominent in formulating evidence to determine probable signals of pollen taxa. Davis and Anderson (1987) collected from modern middens to determine how incorporated pollen reflects the local vegetation and found that the midden pollen was generally representative of the local vegetation. Insect pollinated species were over-represented and these taxa were found in the midden components.

Lower percentages of regional pollen in middens versus other sediments in semi-arid environments, may reflect a lesser importance of air-borne transport for middens. This may be a result of slower accumulation rates for regional pollen or faster accumulation of local taxa (Davis and Anderson 1987). The windblown pollen may express a regional signal although source areas may be variable and windblown pollen can accumulate in the outer rind of middens during periods of high humidity (Thompson 1985). Pollen on plant surfaces incorporated into middens may carry a regional signal but this is biased towards the plant species on which the pollen is preserved (Davis and Anderson 1987).

The issue of over-represented taxa has been observed in middens as there is a tendency for some pollen types to be represented at disproportionate levels to their occurrence in the vegetation. Problems caused by selective over-representation of some taxa might be solved by excluding them from the pollen sum. It is easy to identify an over representation of herbaceous insect pollinated plants but more difficult to decide if wind pollinated shrubs and trees are over-represented. Selectivity by packrats may be the reason for so much "noise" in pollen signals from the middens (Thompson 1985).

Fall et al. (1990) considered that homogenous frequencies of the same pollen taxa and heterogenous of others may indicate different means of pollen deposition and reflect differences in the vegetation from varying distances from the nest. A consistent percentage of background pollen reflecting extra-local and regional vegetation showed assemblages were less biased by rat collecting behaviour. However, this may not always be consistent and at times it is difficult to separate vegetation change from collecting behaviour. Yet, given that packrats are still extant in the United States with a well documented collecting behaviour (within 30-100 metres of the nest site) (Wells 1976; Cole 1990; Spaulding 1990; see studies in Betancourt et al. 1990b; Van Devender and Wiens 1993; ) the role of packrat foraging is becoming less important in more recent research on midden pollen analysis.

There are instances where pollen taxa occur in middens but not in the macrofossil equivalents. Some percentages of pollen are so high that incorporation of flowers into the midden is implied (flowering of some taxa such as Apiaceae and Cruciferae where -17-

plants have flowers and mature fruits in the same inflorescence) or are too high to reflect plant abundances in the local vegetation (eg Cupressaceae, Cruciferae) (Thompson 1985 cited in Davis and Anderson 1986, Davis and Anderson 1987). If macrofossils of a plant are in the midden then its pollen should be considered local irrespective of whether the taxon is known to be wind pollinated and transported long distances (Van Devender 1988).

Plants identified in macrofossil assemblages, but not in the pollen, provide some insights into the limitations of midden pollen analysis. The absence of pollen could be due to; small pollen grains, production of small quantities of pollen with large grains dispersed by insects, grains that are unidentifiable or poorly preserved (Anderson and Van Devender 1991). There is also the factor of seasonal bias in the spectra as it is proposed that the pollen content of packrat middens reflects spring when pollen production is at a maximum (O'Rourke 1991). Anderson and Van Devender (1991) found that the pollen assemblages and macrofossils in middens provided a general view of flora indicating a bias towards summer. Macrofossils also indicated annual flora as their taxonomic resolution allowed identification of summer and winter species. In the Australian context, this is problematic because it is not possible in most instances to identify pollen to species level nor is the flowering so predictable in areas of high rainfall variability.

Macrofossil and Pollen Records Plant fossils and pollen spectra are not directly comparable because of biases inherent in the deposition of each, however palaeoecological reconstructions can be compared (O'Rourke and Mead 1985; Wigand and Nowak 1992; Anderson and Van Devender 1991, 1995). Macrofossils have been reported both quantitatively and qualitatively whereas pollen sums are quantitative. The quantification of macrofossils can vary. They are commonly reported as either raw weights of each group of species of macrofossils, percent weight of selected taxa or the percent weight of the total plant fragments recovered. Ranking macrofossil abundance on a qualitative scale can also be done where 1 = rare and 5 = abundant, allowing records from different midden samples to be readily comparable (Mehringer and Wigand 1990; Spaulding et al. 1990; VanDevender 1990).

Thompson (1985) used a macrofossil: pollen index to compare the two different fossil assemblages. This allowed an examination of how the different data sets approximate each other, and provided a better understanding of the relationship between pollen and macrofossils and the relationship of pollen to the vegetation (Van Devender 1988; Anderson and Van Devender 1991). The pollen is scaled (in relative %) to approximate the relative abundance scale used for plant macrofossils. Theoretically the index should -18-

vary from -5 (extremely abundant in the pollen and not macrofossils) to +5 (extremely abundant in the macrofossils and not the pollen). Each pollen taxon is converted to a scale from 1 to 5 as follows: 5= >50%, 4= 25 - 49%, 3= 10 - 24%, 2= 1 - 9%, 1= <1%, 0= 0%

The index is calculated by "...summing the differences between the relative abundances of macrofossil and pollen for each taxon from all middens in which that taxon was represented by either data type and dividing by the number of samples..." (Anderson and Van Devender 1991:17).

Thompson (1985) acknowledged that macrofossil and pollen assemblages from middens convey the same semi-qualitative signal, however systematic differences occur and include; a reflection in the differences of source areas (extra-local and local vegetation represented in the pollen and macrofossils representing the local plants); the difficulty of assessing whether midden macrofossils and pollen operate on the same time scale although it seems probable that they represent short intervals; and macrofossils provide greater taxonomic precision whereas pollen provides information on taxa that may not be recorded at the site.

Hyrax middens from South Africa indicated that plant material may be absent from middens or, if present, not necessarily collected by the Hyrax (Scott 1990). Macrofossils may have a questionable origin and may not be from the same environment as where the midden was constructed and can lead to spurious interpretations of fossil material.

2.2.5 Australian Midden Studies

There are two known species of stick-nest rat, Leporillus apicalis (Lesser Stick-nest rat) and Leporillus conditor (Greater Stick-nest rat), that are native to Australia. The Lesser Stick-nest rat is now thought to be extinct on the mainland. It was known to have occupied semi-arid to arid shrublands in rocky breakaway country throughout central and Western Australia. The only surviving colonies of the Greater Stick-nest rat are on Franklin Island in the Great Australian Bight and in captive breeding colonies (Copley 1988). The preferred habitat of the Greater Stick-nest rat when extant on the mainland, was low shrubland communities on the open plains.

It is important to understand nest/midden building habits and behaviour of Leporillus spp., as this behaviour partly determines what material is introduced into the nest and consequently what palaeoecological evidence is preserved. However, our ability to do this is confined to historical sources because the rats are extinct in the Flinders Ranges. -19-

Dietary preferences of Leporillus spp. included leaves and fruits of succulent shrub species with high water content (Watts and Aslin 1981; Copley 1988). Newsome and Corbett (1975) found that vegetation preferred by L. conditor included succulent chenopods and Asteraceae species. Grass and seeds were thought to represent a small component of the diet (Watts and Aslin 1981). Work by Read (1984) and Copley (1988) suggests that the diet varied with different seasons, a factor that needs to be taken into consideration when interpreting palaeoecological material from the middens.

Tunbridge (1991) suggests that the rats became extinct on the mainland within 30-50 years of European settlement as a result of pastoral activities causing overgrazing and altered burning regimes. Secondly there was the impact of introduced animals including the fox, cat, goat and rabbit that destroyed natural habitat and/or hunted the rats for food (Newsome and Corbett 1975). According to Copley (1988), the impact from feral animals was more prominent in the northern and western limits of Leporillus' former distributions where pastoralism was absent or took effect later in time.

Work in the field of midden analysis within Australian has been sensitive to methodological issues since the first analysis of Leporillus spp. middens in 1983. Eight studies on Leporillus spp. (stick-nest rat) material have been conducted. These include Green et a/, 's (1983) analysis of a stick-nest rat midden from Gnalta Station in Western New South Wales, Nelson et al. (1990) from the Finke River Gorge in the Northern Territory, Pearson and Dodson (1993) from middens in Western Australia, Berry (1991) from middens in Central Australia, McCarthy et al (1996) in the northern Flinders Ranges, Allen (1996) from the Gap and Coturaundee Ranges western New South Wales and Pearson (1998) on middens from northwestern South Australia and the Northern Territory. These previous studies have served to establish the potential of middens as sources of palaeoecological information and established directions for further work. Overseas work on midden analysis has alerted Australian researchers to methodological issues that need to be resolved and as a consequence, this has become a dominant theme developed in this field.

Green et al's. (1983) study was encouraged by recognition of a similarity between stick- nest rat middens and packrat middens in North America. The palaeoecological potential of stick-nest rat middens was demonstrated by an analysis of the pollen and a chemical analysis of midden contents (Green et al, 1983). The deposit contained pollen, charcoal and large plant fragments. The midden pollen revealed a mix of regional and local signals most likely derived from vegetation found within several kilometres of the midden site (except Callitris), with pollen that originated from a wind blown source and a component derived from the rats' food. This study realised the need for macrofossil -20-

and pellet analysis to further investigate source areas of pollen signals and stratigraphic dating on this type of deposit (Green et al 1983).

Preliminary analysis of a midden from the Finke Gorge National Park in the Northern Territory was undertaken by Nelson et al. (1990). The midden was sampled, divided into three stratigraphic layers and the macrofossils recovered. Macrofossils were identified by matching epidermal patterning of leaves, characteristic venation, stomatal size and pattern and colour with type specimens. Nelson et al. (1990) found differences between the macrofossil assemblages and composition of local present day vegetation. There were approximately 100 species within a few hundred metres of the midden and 18 of these were in the macrofossil record. Five species were recorded in the midden but not the vegetation and both observations were attributed to local fire effects and moisture conditions. The analysis revealed that there had been little change in the vegetation at this location over the last 2 800 years (Nelson et al. 1990).

Berry (1991) analysed plant macrofossil assemblages from two animal middens collected from Kathleen Springs (WSW of Alice Springs) and Mount Swan (NE of Alice Springs) in the Northern Territory. There was an investigation into the progressive change of macrofossil assemblages throughout the middens over a time scale of 3 500 years BP. to 1 700 years BP. No change was detected but there was a weak tendency for the species assemblage to be similar in subsamples from the same layer within a midden. The midden contents were different between sites and reflected the present day geographic variation in vegetation. However no recent middens had been sampled to provide a comparison between the species assemblages that may have been present in middens and the modern vegetation (Berry 1991). This work recognised the geographical constraints imposed by the nature of middens as their preservation depends upon suitable conditions. There was also consideration of the mode of midden construction and the time represented by the accumulation of material This was seen to complicate the interpretation of midden vegetation relationships as there were no currently accumulating middens available to investigate depositional histories (Berry 1991).

Pearson and Dodson (1993) examined the contents of stick-nest rat middens to determine their suitability for providing palaeoecological data from the arid zone. The middens were located in the south western (Bungalbin Hill) and north western (Young Range) edges of Leporillus spp. distributions in Western Australia. They recognised that existing techniques in palaeoenvironmental work lacked sensitivity and a comprehensive geographical spread of sites. Using midden material provided a framework for Holocene data from the arid zone where preservation of this material is not common. Pollen from faecal pellets, grass matting and amberat in the middens were -21-

analysed. These records were then compared to establish whether there was any variation in pollen taxa over time and how representative of the surrounding environment the records were. The pollen record was found to be sensitive to the composition of local vegetation present when the middens were built. There was less wooded vegetation between 900 and 300 years BP with a decline in woody taxa such as Myrtaceae and Dodonaea and increase in herbaceous taxa such as Boraginaceae and Solanaceae.

McCarthy et al. (1996) analysed eight middens from the northern Flinders Ranges. Pollen and macrofossil assemblages were used to reconstruct a discontinuous vegetation record for the Holocene. There were woodlands with a grass dominated understorey in the Early Holocene, shifting in the Late Holocene to vegetation similar to present day communities of shrublands with an understorey dominated by chenopods and grasses. Midden pollen was a mix of local and regional sources and macrofossil taxa were comparable to the pollen assemblages (McCarthy et al. 1996). This research emphasised the need for more detailed dating of midden components to investigate depositional histories of middens. There may be phases of deposition that have occurred which were not detected in the initial sampling. More samples from the Pleistocene/Holocene boundary are also required to test vegetation responses to shifting climatic variables such as the summer/winter rainfall boundary. It is also desirable to increase the number of middens for investigation into the effects of temporal and spatial variability in palaeoecological records.

Analysis of Leporillus spp. middens from the Gap and Coturaundee Ranges, Western New South Wales, examined vegetation change over a ranges of temporal and spatial scales. There was also rigorous dating carried out to investigate accumulation of midden deposits (Allen 1996). It was concluded that deposition of middens is sporadic, though can occur over long periods of time up to thousands of years. Contamination was apparent in some samples and this was regarded by Allen (1996) as a major limitation for palaeoecological interpretations using this type of record. The middens were Mid-Late Holocene age and pollen records suggested a trend from more wooded vegetation to chenopod dominated shrubland (Allen 1996). Macrofossil assemblages were fragmentary but thought to be complementary to the pollen record. The overall consensus was that middens are useful for examination of vegetation and faunal change however the evidence should be treated cautiously and regarded as separate rather than continuous records over a stratigraphically defined timescale (Allen 1996). Discontinuous deposition of the material increases the generality of palaeoecological interpretations. -22-

Pollen and macrofossil records from Leporillus middens less than 3 500 years old suggest little evidence for climatic change during the Late Holocene from central Australia as vegetation communities were stable (Pearson 1998). Radiocarbon analysis was used to help develop a conceptual model of accumulation of middens to understand the taphonomy of these deposits. Pearson (1998) concluded that analysis of middens are currently the best source of palaeo-environmental information in the arid zone of Australia.

This current project builds on from the previous research by including a larger number of middens sampled from a more extensive geographical area in the Flinders Ranges. Detailed dating of individual middens and separate components (i.e. leaves, faecal pellets and pollen) will allow an investigation into intra-midden variability and increase our understanding of taphonomic processes. New palaeoenvironmental records from the semi-arid zone will contribute to gaps in our inderstanding of the Holocene as discussed in the following section.

2.3 Holocene Palaeoenvironments of Australia

This section discusses Holocene palaeoenvironments of Australia that have been inferred from lake levels, pollen sequences from bogs/swamps, dune and fluvial records. Different lines of evidence used for reconstructing Quaternary environments vary in accuracy and geographical and temporal scale. In various combinations, these lines of evidence indicate periods of aridity, long versus short term change, catastrophic versus gradual change, human versus climate and ecology versus climate change during the Holocene. Pollen core data and lake levels form the basis for the reconstruction of climates (Kershaw 1995) as changes in lake levels and salinity are associated with changes in the regional vegetation cover (Crowley and Kershaw 1994). Most information only allows interpretation in terms of the effective precipitation, not the separation of temperature and actual precipitation or the effects of seasonality (Llyod and Kershaw 1997). However records from some sites (discussed in this section) argue evidence for detecting seasonal shifts in summer and winter rainfall regimes and the influence of ENSO causing variability in climate during the Holocene.

There is regional variation in Holocene climatic change across the continent (Ross et al. 1992) and a clear understanding of regional patterns is limited by the patchy distribution of records and bias to well watered areas of the continent (Thorn 1992; Kershaw 1995; D'Costa and Kershaw 1997), thus emphasising the importance of semi- arid palaeoecological records. -23-

2.3.1 Overview of Holocene Climate

The Pleistocene-Holocene transition was characterised by dramatic environmental change ranging from the height of the Last Glacial Period with sea level 120 metres lower than present and 30% of the earth covered in ice sheets, to attainment of present day conditions approximately 6 000 years BP. (Kershaw 1995). However, it is important to note that there appears to have been regional variation in rates and the timing of amelioration (Kershaw 1995). Interglacial periods account for approximately 10-15% of the Quaternary, suggesting that species and communities present at these times may be atypical (Hope 1994).

At 10 000 years BP. there was a rapid climate change observed in most regional records in Australia, with temperature and precipitation reaching close to present day conditions. This was reflected in the pollen records with a marked change in assemblages (Kershaw 1995). With increasing rainfall in the north and a decrease in temperature in the south, DeDeckker et al. (1988) suggested high summer and lower winter rainfall at this time.

For most parts of Australia, records suggest that between 8 000-7 000 years B.P, temperatures had risen and the climate was characterised by warmer and moister conditions than present, with most lakes full (Bowler et al. 1976; Chappell 1983; Kershaw 1989; Ross et al. 1992; Nanson et al. 1992; McCarthy et al. 1996). Alpine studies in New Guinea, Mt Kosciusko and Tasmania (Macphail 1979; Hope 1986; Kershaw et al 1986; Markgraf etal.l9$6; Kershaw and Strickland 1990) show warming conditions and invasion of alpine tundra by herb fields after 13 000-14 000 years BP. and temperatures of the present established by 9 500 years BP. In the temperate zone of Australia, maximum warming and moisture was attained between 8 000-6 000 years BP. with rainfall 5%-10% and temperature 1-2°C higher than present. Associated with these conditions was a denser vegetation cover (Kershaw 1995; Lloyd and Kershaw 1997). In central Australia, there were higher levels of effective precipitation and denser vegetation cover between 8 000 - 4 000 years BP (Ross et al. 1992). Increase in precipitation into the Middle Holocene for Southern Australia is a result of a shift in the southern margin of the subtropical high pressure belt north of its present position (Kershaw 1995) and an increase in the influence of westerly winds (Harrison and Dodson 1993). This pattern would be intensified by a decrease in seasonality, rising sea levels and warming ocean surface waters (Kershaw 1995).

The Late Holocene was drier and cooler (Ross et al 1992). Since 5 000 years BP, there has been a contraction in humid habitats and small cooling or increased seasonality has taken place (Dodson et al 1992). The timing of linear dune building Australia wide -24-

includes a late Holocene phase around 4 000 - 1 000 BP (Wasson 1989). This episode could be attributed to increased wind speeds, frequency of drought and Aboriginal burning. The phase corresponds to the period of cooler drier conditions, post dating the high lake levels in the Mid-Holocene (Ross et al. 1992).

2.3.2 Lake Level and Pollen Records

Lakes in Australia have yielded rich records of environmental change (Thorn 1992) as they reflect changing levels of precipitation and evaporation (Ross et al. 1992) The late Quaternary history of the volcanic plains in Victoria, extending into South Australia, is documented from sedimentary and pollen analysis from lakes and swamps (Crowley and Kershaw 1994). Harrison and Dodson (1993) referred to in Kershaw (1995) have provided a summary of the data for the interior of South Eastern Australia. Luly (1990) notes that there was variation in the timing of the rise in levels of different lakes after 10 000 years BP

Generally, there were low levels until 7 000 years BP followed by maximum levels (wetness) around 4 000 years BP and then a gradual drying out to the present. D'Costa et al (1989) demonstrated from pollen and microfossils in lake cores from southwest Victoria, an argument for an increase in precipitation from 10 000-8 000 years BP with moist conditions until 8 000-6 000 years BP This was followed by a decrease in precipitation at 5 000 years BP and also after 3 000 years BP. Lakes from northeastern Queensland (Kershaw 1981) and South Australia suggest a wetter period between 6 500-5 000 years BP (with high lake levels) and falling by 5 000-4 000 years BP while Lake Leake, South Australia, was wet at 8 000-3 500 years BP. (Dodson 1974b).

Lake and swamp records from lower south east South Australia display similar trends in water levels and vegetation during the Holocene. The pollen sequence from Lake Leake indicated dry swamp conditions and woodland with grassy understorey at the Pleistocene-Holocene boundary. Water levels rising into the Middle Holocene corresponded with a major expansion of wet heath vegetation dominated by Casuarina. Precipitation levels were rising faster than evaporation induced by rising temperatures. Low lake levels in the Late Holocene induced swamp conditions and increases in Cyperaceae and Restionaceae plants, and wet heath communities replaced by sand heath. A decline in trees and shrubs and increase in herbs, grasses and Pinus characterised Post-European vegetation communities (Dodson 1974a; 1975). Wyrie Swamp, lower south east South Australia showed similiar trends with Pleistocene Eucalyptus woodland replaced by Casuarina dominated wet heath at the Pleistocene- Holocene boundary with shallow permanent water in the swamp. Increasing water levels during the Middle Holocene corresponded with Casuarina dominance until -25-

approximately 7 000 years BP followed by an increase in Eucalypt woodland (Dodson 1977a). Around 7 000 years BP at Mount Burr Swamp, there were Casuarina and Eucalypt woodland communities as water levels in the swamp were increasing. An expansion in wet heath and myrtaceous scrub occurred from 7 000 - 5 000 years BP then Late Holocene periodic wetting and drying phases saw increased dominance of Casuarina and fluctuating abundances of Eucalyptus and myrtaceous scrub (Dodson and Wilson 1975). The pollen sequence from Lashmar's Lagoon on Kangaroo Island (Clark and Lampert 1981; Singh et al. 1981; Clark 1983) indicated a wet Middle Holocene (aproximately 6 000-4 800 years BP), with vegetation dominated by Casuarina woodland. Drier Late Holocene (until approximately 1 300 years BP) conditions, with an increase in the frequency or intensity of fire, saw expansion of Eucalyptus woodland and woody shrubs replacing grasses. Post-European vegetation was characterised by an increase in grasslands with almost complete removal of Casuarina.

At Lake Bolac in Victoria, there were dry conditions at the end of the Pleistocene, and on refilling, the lake became saline. Vegetation became dominated by Allocasuarina verticillata with the increase in water levels. This may have been synchronous with other sites (Crowley and Kershaw 1994). The dominance of Allocasuarina ended at approximately 8 000 - 7 000 years BP (probably due to an increase in the saline ground water) and was replaced by Eucalyptus to form riverine woodland communities. An exact date for this shift in vegetation communities is not definitive due to regional variation in this event (Crowley and Kershaw 1994). Swamp conditions began around 4 000 years BP in response to a decrease in precipitation. During European occupation, there was an observed change in fire patterns, reduced tree cover, loss of floristic diversity, a decrease in lake levels and an increase in salinity (Crowley and Kershaw 1994).

Lake levels in northwestern Victoria, ie. Lake Keilambete, Gnotuk and Bullenmerri were lowest and salinity highest, prior to 8 300 years BP. The lakes began to fill and by 6 500-3 800 years BP, levels were high with lake full stage attained at 6 400-5 700 BP (White 1994). There was a dry phase when levels were low between 3 000-2 000 years, followed by fluctuating levels and an increase in the last 1 000 years but not to previous levels experienced in the Middle Holocene (Bowler 1981; Creagh 1993). Pollen (Dodson 1974a) and ostracods (Chivas et al. 1985) verify this history based on the sedimentology and mineralogy.

Lake Tyrell in western Victoria was shallow at 7 500 years BP and from 6 500-3 200 years BP, levels were higher giving the lake a semi-permanent status. At 2 000 years BP the current playa phase was established (Teller et al. 1982). Luly's (1990) findings -26-

based on lake levels and the pollen record are in general agreement with this previous work. Lake Tyrell levels mesh with other Holocene records from south eastern Australia (Luly 1990).

Luly (1990) emphasises that corresponding changes in Holocene lake levels and geomorphology are detected in pollen studies from arid areas. There were shifts in vegetation communities in response to changing precipitation at Lake Tyrell. Mallee Eucalyptus and Callitris woodlands replaced Casuarinaceae between 6 600 and 2 200 years BP under higher rainfall conditions (Luly 1993). From 2 200-800 years BP mallee persisted and Casuarinaceae expanded but there was a drastic decline in Callitris. Casuarina then dominated the vegetation community forming a more open cover. When sufficient rainfall was available, there was an expansion of grasses (Luly 1990).

Lake Eyre is an ephemeral playa lake that receives most inflow from the northern Australian monsoon and the sedimentary and palaeohydrology form a record of the monsoon runoff (Magee 1997) There is evidence of minor lacustrine events during the Early Holocene, following a dry phase. Establishment of modern ephemeral conditions occurred between 3 000-4 000 years BP (Magee 1997). The lake filled episodically and achieved a semi-permanent status between 10 000-5 000 years BP and the current playa status was established at 5 000 years BP (Ross et al. 1992) that corresponds with Nanson et al's. (1992) claim that the Late Holocene was dry.

A peat deposit in the spring complex at Dalhousie Springs in Central Australia, provides a local history for the last 2 000 years (Boyd 1990; Boyd 1994). Gibber plains surrounding the spring were characterised by chenopods and herbaceous vegetation, whereas perennial shrubs and some trees were found to colonise the floodplains. The pollen was predominantly local in nature (Boyd 1990). There has been an expansion of swamp conditions, indicating a reduced flow and hence less effective precipitation during the Late Holocene (Boyd 1994).

Martin (1973) examined the palynology and historical ecology of cave sediments from the Nullarbor Plain in Western Australia. Holocene age pollen sequences were analysed from two excavations near Madura and a Pleistocene record from Eucla. Myrtaceae and Chenopodiaceae fossil pollen demonstrated a reciprocal relationship. That is, when the Myrtaceae pollen was abundant in the sample, the Chenopodiaceae pollen levels were depressed. Surface pollen samples also displayed similar trends in the Chenopodiaceae/Myrtaceae ratio that reflected relative abundances of the two floristic groups in the modern vegetation (Martin 1973). -27-

At 6 000-5 000 years BP at Eucla there was an increase in mallee scrub, as sea level rose and precipitation levels increased. Unlike Eucla, Madura records showed a marked decline in the mallee scrub cover in the Middle Holocene. This was attributed to an increased frequency of Aboriginal burning in the landscape in an area of relatively lower annual rainfall compared to Eucla. Hence, the mallee regeneration rates were thought to be lower at Madura after fire, but more extensive at Eucla (Martin 1973).

The pollen sequence from N145 rockshelter, Nullabor Plain, South Australia suggested a treeless plain at 10 000 years BP. During the Early and Middle Holocene there was an increase in mallee cover, remaining to the present (Martin 1973).

2.3.3 Dune and Fluvial Records

Alluvial fans on the western side of the Flinders Ranges began to aggrade prior to 5 000 years BP and continued to accumulate until approximately 3 500 years BP after which time the fans were incised as the Middle Holocene phase was followed by an increase in aridity (Williams 1973). There was also accumulation of alluvial fans near Broken Hill, between 6 000-3 000 years BP (Wasson 1979). Dunes in the northern Strzelecki Desert were stable between 8 000 and 4 000 years BP however, dunes adjacent to Balcoracana Creek (south-western end of Lake Frome) were built during the Early to Middle Holocene suggesting a drier climate (Ross et al 1992). Dunes were being formed from approximately 2 500-640 years BP in western New South Wales and then stable and vegetated since 640 years BP (Wasson 1976). Approximately 3 000-2 000 years BP, dune building was occurring at Cobar (Chappell 1991) and there was comparable dune activity identified in the Victorian Mallee district, Simpson-Strzelecki dunefields (Luly 1990; Ross et al. 1992) and in eastern Australia (Cook 1986).

2.3.4 Evidence for Seasonality in Holocene Climates

A water level for Lake Frome from Bowler and Magee (1990) shows evidence for refilling after deflation of the lake floor during the Holocene. Singh and Luly (1991) suggested that grass pollen reflected incursions of summer rainfall into the lake catchment while Asteraceae reflected winter rain. The palaeoclimate in summarised as follows: 14 500-13 000 Low summer and declining winter rainfall: dry period 13 000-6 000 Dominance of summer rainfall; wetter 6 000-4 500 Total rainfall increased; dominance of summer rain 4 500-2 200 Decreased total rainfall; winter rainfall certain/ uncertain summer rainfall 2 200-0 Slight increase in total rainfall as result of an increase in summer rain -28-

According to this interpretation, the increase in lake levels was likely a result of increased run off from summer rainfall. From 9 500-8 000 years BP Eucalypt woodland, shrubland and grasses flourished with enhanced summer rainfall. There was a decline in trees, shrubs, grasses and chenopods and increase in ephemeral taxa from 8 000-7 000. During the wetter Middle Holocene (7 000-4 000 years BP), trees shrubs and grasses recovered while ephemerals declined. This cycle repeated from 4 000-2 200 years BP. During the Late Holocene (2 200 years BP- present), with an increase in the dominance of winter rainfall, chenopods expanded, grasses declined and there was a small increase in trees and shrubs (Singh 1981; Singh and Luly 1991). Vegetation in the Flinders Ranges owes much of its present nature to aridity and the prevalence of winter rain (Gell and Bickford 1996).

2.3.5 Contradictions Between Holocene Climate Reconstructions

Higher temperature and precipitation levels than present during the Middle Holocene are not supported by records from some sites in Australia. Examples from The Great Sandy Desert of northwestern Australia, Rottnest Island southwestern Australia, Fraser Island on the east coast and Groote Eylandt northern Australia will illustrate the contradictions.

At 6 000 years BP from Papua New Guinea to southern New South Wales, the mean annual temperature was between 0.5°C-4°C higher than present and effective precipitation was at its peak. In south-western Australia, climate was different as north of 35° latitude the area was dry and precipitation was 90% of the present levels (Ross et al. 1992:97). Late Holocene (2 000 years BP) showed changes sometimes contradictory between sites, while other sites within Australia recorded no change. Temperatures in Tasmania and southern New South Wales dropped by as much as 3°C and northern Queensland temperatures were about 1.5°C higher than present (Ross et al 1992).

Wyrwoll et al (1986) and Wyrwoll et al (1992) used stratigraphic data from swamps and alluvial sequences to reconstruct Holocene climates from the Northern Great Sandy Desert and concluded that there has been little change in the intensity of the north-west monsoon since approximately 6 500 years BP and that monsoon activity may have been reduced during the Early Holocene. There was no indication from sediments or the pollen record of any change in the flood regime over the last 6 500 years, arguing less surface water during the Early Holocene, thus suggesting a region that was more arid. Taxa such as Cyperaceae and Typhaceae representing swamp communities, were present throughout the sequence. Pollen spectra did not adequately register any change in the regional record as the pollen found in the cores reflected locally occurring species (Wyrwoll et al. 1986). This work is not in agreement with Singh and Luly -29-

(1991) or Wasson and Donnelly (1991) that suggests there have been significant changes in areas affected by the northern Australian monsoon. This is possibly a factor of differences in the resolution of different palaeoecological records.

Reassessment of previous palynological studies from southwest Western Australia by Newsome and Pickett (1993) suggests that conditions in the Middle Holocene were similar to the present or perhaps slightly more arid. From other evidence using coastal geomorphology from Northern Australia, Lees (1992) claims a Holocene drying trend beginning between 5 000-4 000 years BP and finishing at approximately 2 000 years BP. It is important to note that there is regional variation in the timing of climatic change throughout Australia that reflects the present day climatic gradients inland from the coast and north along the coast (Ross et al. 1992).

A pollen record and lithological study of a core from Barker Swamp, Rottnest Island indicated between 7 500-6 600 years BP a fresh open water lake was established and surrounded by sedges, Callitris low forest and restricted jarrah/eucalypt woodland communities were present. It was argued for increased aridity around the Middle Holocene (5 300 years BP) with sparser vegetation with and increase in Asteraceae and Chenopodiaceae pollen and decline in eucalypts (Backhouse 1993).

Longmore (1995; 1997a; 1997b) presents a record from perched lakes on Fraser Island that suggests a Middle Holocene dry period via a reconstruction of past lake levels and shifts in the precipitation/evaporation ratios. The pollen and charcoal evidence suggests a change in the vegetation and burning regime during the Middle Holocene. Casuarinaceae increased from the base of the core (dated to 8 460 years BP) and peaked between 5 500-4 500 years BP and charcoal increased and peaked at 5 500 years and 3 000 years BP. At 2 500 years BP, both levels had fallen. Rainforest taxa increased by 1 500 years BP (Longmore 1995). In summary, there was a fall in the water table and increased period of burning suggesting a drier period during the Middle Holocene, followed by recovery of the lake level and expansion of rainforest components in the vegetation (Longmore 1995).

In the earliest part of the Holocene, northern Australia was comparatively arid and the timing of the Middle Holocene maximum was approximately 1 000 years later than that for southern Australia based upon pollen evidence from Groote Eylandt (Shulmeister and Lees 1995). The work summarised in White (1994) shows that the pollen sequence from Four Mile Billabong, a perennial lake in the northeast of the island, indicates there was permanent water in the lake by 9 000 BP with a swamp community of Melaleuca, water lilies and Myriophyllum (Haloragaceae). The surrounding dunefield was open grassland. From 9 200-7 500 BP paperbark was still present and then to 5 000 years BP -30-

open forest replaced grassland with Eucalyptus and Acacia shrub. Between 5 000 and 3 800, lake levels were high. Levels then fluctuated up until 1 000 years BP where grassland expanded with the increase in occurrence of cool fires (White 1994). Climatic fluctuations were reflected in the pollen in the Late Holocene with a decline in effective precipitation between 3 700-1 000 years BP followed by a recovery to modern values (Schulmeister and Lees 1995). It is suggested that ENSO-induced climatic variability is a feature of the last 3 000-5 000 years and the Early Holocene was less variable (McGlone et al 1992; Shulmeister and Lees 1995). A combination of geomorphic and pollen records strongly support a dynamic climate over the last 3 700 years for northern Australia with: 1) continuous increase in effective precipitation from 10 000-5 000 years BP; 2) Maximum effective precipitation between 5 000-4 000 years BP; 3) Decline in effective precipitation from 4 000-3 500 years BP; 4) Recovery of effective precipitation for less than 2 000 years BP (Shulmeister and Lees 1995).

Contradictions between different records are occurring because of factors such as; a) lag time in responses particularly with pollen data and vegetation records b) continental variability when comparing records from south-east, north and south­ west regions in Australia c) resolution of different data sources for example, where geomorphic evidence is broad scale in nature in contrast to high resolution pollen data from lake cores d) thresholds of change e) complicating issues such as the role of fire and human impacts

2.4 Contribution of Leporillus spp. Midden Records

Hall (1986) suggested that tight spatial resolution of middens indicates they are unsuitable for regional palaeoenvironmental reconstructions, because the records are local. However Spaulding (1990) countered this by stating that knowledge of the source area is an advantage in the field of palaeoecology. Different vegetation chronologies at sites that are less than 200 km apart reflect regional vegetation change throughout the Pleistocene and Holocene. Increasing the number of sites may allow the recognition of a mosaic pattern of late Quaternary vegetation change whereas a temporal series of midden pollen spectra can reflect the large scale vegetation change that has been detected in stratigraphic pollen records and macrofossil assemblages (Thompson 1985). These principles can be applied to Leporillus spp. midden studies. Records from Leporillus spp. middens make an important contribution to the resolution of palaeoecological evidence for the semi-arid zone. Middens offer a high spatial resolution as they are located throughout an extensive geographical region that includes the southern arid and semi-arid zones of the Australian continent. This location is likely to be sensitive to changes in an important palaeoclimatic parameter, the summer -31-

rainfall/winter rainfall boundary. There is good temporal resolution as individual middens provide local and regional scale information on vegetation patterns over time. Multiple sources of palaeoenvironmental evidence (ie. pollen and macrofossils) from a single deposit is also a major benefit. There is excellent preservation of pollen, while preservation and abundance of macrofossils is more variable. Finally, the middens provide a methodological comparison with the pollen record from Lake Frome (pollen core compared to the more diverse range of plant evidence found in middens) and a biogeographic comparison (arid playa lake compared with adjacent mountain range). The latter can provide information on the palaeoecology of refugia within the semi-arid zone.

2.5 Conclusion

Research from overseas and within Australia has shown that the potential and application of midden analysis has been successful while at the same time drawing attention to the need for further investigation into methodological and taphonomic issues. Macrofossil analysts should include pollen analysis as a standard analytical procedure in midden analysis (Van Devender 1988). Pollen in middens is abundant, well preserved and complements the macrofossil analysis because it reflects local and regional vegetation patterns. Comparison of pollen and macrofossils will help evaluate the sensitivity, accuracy and utility of midden pollen and may provide insight into the presence of a species much older than may appear in the macrofossil records (Van Devender 1988). It must be recognised that variability among the middens can be high and the accuracy with which pollen reflects the vegetation will vary (Davis and Anderson 1987).

Variability in pollen assemblages from different midden samples can be attributed to environmental site differences, recent vegetation change and midden characteristics. Local and regional vegetation patterns can be detected from the records (Davis and Anderson 1986) but there can be difficulty in interpreting the records, caused by uncertainties in recognising pollen source areas (O'Rourke and Mead 1985; Davis and Anderson 1991). The validity has to be established through a systematic collection of modern middens and comparing vegetation data from different vegetation types (Davis and Anderson 1986). A combination of regional and local pollen signals and macrofossils preserved in middens provides a more accurate picture of palaeovegetation communities (King and VanDevender 1977; Cole 1990).

The regional record from packrat middens requires many dates from many middens collected at different sites. Midden sites including canyons and rocky uplands may be regarded as atypical, imposing a microclimate which impacts on the wider vegetation -32-

(O'Rourke and Mead 1985). This factor must be acknowledged as a limitation in midden analysis (Spaulding 1990).

Historically, the use of middens for palaeoecological information in arid areas has been used as a basis for reconstructing the palaeoecology of the south-western USA. As a consequence of the pioneering work from the United States, a number of methodological issues that require investigation have been discussed to an extent in the literature. The most important of these are 1) developing an understanding of depositional histories of this type of deposit and 2) investigating source areas and recruitment processes of pollen preserved in the middens. However, given the long term records available from Neotoma middens and the clear zonation of vegetation communities at different altitudes, the focus of research has been on the reconstruction of long term records using plant macrofossils. Pollen is increasingly included is palaeoecological reconstructions using midden records because of the complementary role to macrofossil inventories (Anderson and Van Devender 1991; Anderson and Van Devender 1995) and 'traditional' sedimentary deposits for pollen analysis are rare within lowland arid regions in the US (Anderson and Van Devender 1995). Studies have also been conducted whereby macrofossil records from middens are compared to pollen records from traditional sediment/lake cores.

In other countries, including Australia, the emphasis has been placed on pollen midden analysis and resolving methodological issues that improve integrity of palaeoecological interpretations. Plant macrofossils are less abundantly preserved in these middens and the pollen record is relied upon in more instances. Characteristics of vegetation communities also differ from the US studies, with a less distinctive influence from the altitudinal zonation of plant species that is clearly recorded in Neotoma middens.

Records from Leporillus middens makes an important contribution to gaps in the knowledge of Holocene palaeoenvironments, based on the high spatial resolution and good temporal resolution of pollen and plant macrofossil records.

The following two chapters illustrate where and how my research was undertaken. The regional setting where stick-nest rat middens have been sampled will set the scene, followed by an outline of the sampling strategies used in the field and laboratory, to investigate methodological and taphonomic issues in analyses of stick-nest rat middens. -33-

Chapter 3: The Flinders Ranges: A Regional Perspective

3.1 Introduction

This chapter describes the regional setting of the Flinders Ranges with an overview of the geology, climate and the nature of semi-arid vegetation communities. This leads into a description of individual midden sites within the context of their environmental settings and associated vegetation communities. The spatial variability between each study site, in terms of the location within the northern and central Flinders Ranges and proximity to the summer-winter rainfall boundary, is addressed. This has direct implications for the interpretation of pollen and macrofossils from the middens. Consideration of the Flinders Ranges as a refuge will be discussed as it is important to establish the sensitivity of this environment's response to climatic and other ecological change and to be aware of this for palaeoecological and palaeoclimatic reconstructions for the region. The chapter concludes with the implications of the regional context for the themes of this research project.

3.2 The Flinders Ranges

The Flinders Ranges, approximately 62 000 square kilometres in area, are located in the semi-arid zone to the north east of Spencer Gulf in South Australia (Greenwood et al. 1989) and extend from the Mount Lofty Ranges in the south to the Strzelecki dune fields and Lake Eyre Basin in the north (Symon 1972; Lampert and Hughes 1987) (Figure 3.1). The southern margin of the ranges is 300km north of Adelaide and extends another 400km to the north (Domin 1986; Cock et al. in press). It is a ragged landscape with much folded, faulted and uplifted Pre Cambrian and Palaeozoic massive beds of sandstones interleaved with softer shales that have been differentially eroded (Fox 1991; Lemon 1996; Mincham 1986). There are a variety of topographic environments including high plateaus, ridges, gorges, cliffs, boulder slopes, terraces, tablelands, valleys and plains (Corbett 1980; Cook 1986b; Lampert and Hughes 1987; Fox 1991). This diverse landscape rises from below sea level at Lake Eyre to 1 170 metres above sea level at St Mary Peak in Wilpena Pound (Swinbourne 1986).

3.2.1 Geological Setting

Regional geology is a mix of quartzites, granites, slates, limestones, dolomites, sandstones and shales (Fox 1991). Ancient rock beds originated as marine deposits in a long trough and, on the subsiding sea floor, sand and silt accumulated to depths greater than 20km. After subsidence, crustal forces compressed, buckled and folded the sediments, and along lateral lines of weakness, uplifted rock beds to be eroded -34-

Ploya loke A y/\ Ranges • Settlement

0 100 i_ —I kilometres

o Loke Eyre r O

Marree

fyArkaroola

Copley^ Z.oAre i Loke *> Frome •31' U) Torrens

Port Augusta

EYRE 1 PENINSULA * 4 £-N / ot "•rx^ « "f^ 1 CyM>"'/ |'"N. V^ V- SPENCER \"- sJM'C \pJ i GULF fo' . i/ADELAIDJ/' / E l 1 —< 4v*J/^ 1 Ci

Kangaroo Island 139* _J

Figure 3.1: Regional location map of the Flinders Ranges in South Australia. -35-

(Mincham 1986). The oldest rocks (Palaeoproterozoic to Cambrian) are found in the northern Flinders Ranges and include the Freeling Heights Quartzite in the Mount Painter Complex. These are highly metamorphosed and heavily dissected (Selby 1990). There are mineralised rocks underlying the sedimentary rocks that have been uplifted and exposed, forming the only granite country in the Flinders Ranges (Mincham 1986). Between Copley and Arkaroola there are a series of folded sequences that represent shallow marine depositional environments and include the Tapley Hill Formation and Amberoona Formation (laminated green siltstone) of the Umberatana Group and the overlying Nuccaleena Formation (dolomitic shale), Brachina Formation (green and purple siltstone), Bunyeroo Formation (red-brown shale), Wonoka Formation (green shale and limestone) and Pound Quartzite forming the Wilpena Group. Sandstones and shale outcrops form the lower hills, ridges and valleys (Twidale 1980; Selby 1990).

3.2.2 Climate

The Flinders Ranges are in the driest state of Australia (Swinbourne 1986). Regional rainfall is low and highly variable from year to year (Cock et al in press). There is an increase in the incidence of summer thunderstorms across the Ranges compared to the plains, as the central mountains produce an orographic effect. According to Schwerdtfeger and Curran (1996), orographic influences increase during winter months as falling condensation levels intercept the higher topography, whereas in summer, the condensation level rises to higher altitude and intercepts less topography. Lampert and Hughes (1987) report that the northern ranges receive an annual rainfall of just less than 300 mm, which is 50% greater than the adjacent plains (125 mm on Lake Frome), while the central ranges (Brachina Gorge) receive an average of 200mm per year and an average annual evaporation of >2000mm (Cock et al in press). Medlin (1993) has reported < 150mm per year for the eastern central ranges (Mount Chambers Gorge) with an average evaporation rate of approximately 2700mm per year, averaging 90mm in July and 400mm in January. The summer/winter rainfall boundary (Figure 3.2) crosses the continent to the north of the ranges (Singh and Luly 1991). Seasonal variations in the climate result from the shift in the position of the high pressure belt from the southern region of the continent in summer to central Australia during the winter (Winkworth and Thomas 1974; Allan 1990). During winter and spring, low pressure systems with their associated cold fronts, bring rain to southern South Australia and rain falls in the northern and central Flinders Ranges only if the cold fronts penetrate far enough into the northern part of the State. Blocking highs centred over Tasmania may sometimes deflect the lows, resulting in lower than normal winter rainfall (Medlin 1993). The high and low pressure systems shift south during summer and autumn and the cold fronts rarely extend inland. There is an incursion of tropical systems from -36-

Figure 3.2 Location of the summer rainfall/winter rainfall boundary in Australia and proximity to Lake Frome (after Singh 1981: 421). -37-

northern Australia where tropical lows provide summer rain. These rains may penetrate deep into South Australia below the summer/winter rainfall boundary (Medlin 1993).

Daily maximum temperatures usually range between 18°C (winter months) and 35°C (summer months) in the northern Flinders Ranges (Sprigg 1984) and 20°C (July) and 34°C (January) in the eastern central ranges (Medlin 1993).

3.3 The Semi-Arid Environment: Factors Influencing Vegetation Communities

Characteristics of semi-arid vegetation communities are a result of environmental factors that include rainfall/climate regimes, physiographic features and the role of disturbance. These aspects that affect the composition and structure of vegetation communities will be discussed, to serve as an explanation of the variable nature of semi-arid vegetation at a regional level. It can then be demonstrated that the combination of these factors at a regional scale are responsible for the observed diversity of vegetation communities recorded in the Flinders Ranges.

3.3.1 Physical Environment

Physiographic factors such as topography and soil type affect the character of vegetation communities over a wide range of geographical scales. According to Mabbutt (1984), physiographic differences fix limits on broad vegetation types characteristic of the arid and semi-arid zones, because climatic boundaries are essentially transitional. Topographic diversity and moisture accumulation associated with rocky environments results in a greater diversity of species (Freeland et al 1988).

A generalisation has been made that desert uplands support a variety of plant species due to the wide range of microhabitats for plant communities to colonise (Mabbutt, 1984). In the Flinders Ranges, variable geology and topographic settings lead to high spatial variability in the vegetation. Shaded rock clefts trap moisture and debris providing suitable conditions for the growth of deep rooted trees and shrubs. Debris slopes have little infiltration and this is ideal for xeric spinifex communities. Trees and large shrubs grow on the lower slopes receiving runoff from the higher slopes on a mantle of loose sediment.

3.3.2 Influence of Rainfall

Rainfall is another factor that influences the structure, composition and distribution of species of flora in semi-arid vegetation communities (Carrodus and Specht 1965, Mabbutt 1984). It is important to know the seasonal distribution in addition to the total amount of rainfall to understand and interpret vegetation communities (Walter 1971). -38-

Highly episodic rainfall controls the flux of perennial plant populations (Stanley, 1983) as low precipitation will inhibit pollen production (Green et al. 1988). A change in the distribution of perennial vegetation from an abundant cover to a contracted distribution is a response to water deficiency (Carrodus and Specht, 1965). When there is adequate raninfall during the year, the vegetation regenerates vigorously (Walter 1971). Ephemeral species depend on rainfall at any time of the year and are referred to as buffering vegetation as they have a range of growth rates and water requirements (Walter 1971). Annual species only germinate within suitable temperature regimes (Walter 1971) and there can be variation in the population size of annuals in the case of drought periods (Green et al. 1988; Fox 1991). Therefore annuals may not be present from year to year. Generally after winter rains, semi-arid vegetation communities are dominated by Asteraceae spp. and after summer rains, Poaceae spp. and Amaranthaceae spp. flourish.

Knowledge of wet and dry periods and their duration is important in understanding the growth and decline of plant species in the Flinders Ranges (Medlin, 1993). There is winter rain in the south and summer rain in the north and the rainfall in between these regions can be unpredictable (Walter 1971). The location of sites under the influences of both summer and winter rainfall regimes contributes to a higher species richness in vegetation communities (Keighery and Gibson 1993). Mount Chambers Gorge and Brachina Gorge experience summer and winter rainfall while the Arkaroola-Mount Painter Sanctuary experience summer rainfall and a weaker influence from winter rain. Medlin (1993) suggests that good summer rains followed by winter rains for three to four years in succession are required for the regeneration of many plant species. However, rainfall variation can introduce a problem in the perception of vegetation change as extremes of high and low events and/or the co occurrence of two rare events (eg. fire and an extreme wet period) can lead to occasional shifts in species densities that can persist for years (Noble 1986). Short term change in rainfall can affect ecological changes and short term low amplitude oscillations in the mean annual rainfall are reflected in the pollen record (Green et al. 1988). These are all factors to consider when interpreting pollen and plant macrofossil records from the semi-arid zone.

3.3.3 The Role of Disturbance

Grazing lands are predominantly found in semi-arid or sub-tropical zones with extremes of climate or terrain and where pastures or intensive agriculture cannot be sustained (Wilson 1990). However, arid zone vegetation is poorly adapted to continuous grazing (White 1994). Pre-European vegetation had elements of high quality ephemerals, medium quality perennials and palatable species (Harrington et al 1984), however the -39-

introduction of domestic stock has caused a major change in the structure of native vegetation (Cunningham et al. 1992). There has been a decrease in perennial herbs, forbs and grasses (nutritious forage in the dry season) and increases in annual herbaceous and ephemeral plants, inedible woody plants, shrub density (due to reduced competition from perennial grasses and changes in fire regime), as well as soil erosion and a redistribution of nutrients (Harris 1983; Low 1984; Hobbs and Hopkins 1990; Wilson 1990; Mitchell 1991; Cunningham et al. 1992; White 1994). Grazing pressure on ephemerals is less than perennials because brief periods of rapid growth exceed their consumption by stock.

In the Flinders Ranges in the early 1860s, favourable seasons lead to an over estimation of the stock carrying capacity of the land and in the mid 1860s in the Great Drought, vegetation was eaten bare and stock died in the thousands (Mincham 1986). The long term response of plant communities to grazing depends upon the intensity of defoliation and the sensitivity of each species. Tolerance to defoliation varies between species. For example, Atriplex vesicarius is a sensitive species that may not regenerate whereas grasses vary in their tolerance. Some Themeda spp. are classed as sensitive, Enteropogon spp. fail to set seed if defoliated when young and Aristida spp. are resistant and regarded as prolific seeders in grazed lands (Wilson 1990). Chenopods are also not fire tolerant and when part of an understorey in woodlands where fire is a greater risk, they tend to be replaced by grasses and woody shrubs (White 1994).

Chenopod shrublands have adapted to survive erratic rainfall, extreme temperatures, soil salinity and grazing by native animals but not the effects of hoofed grazers or rabbits (Williams and Oxley 1979; Pickard 1991). Atriplex vesicarius is most palatable to stock. Grasses and forbs that grow between the chenopods constitute much of the fodder when there has been rain however when they are gone, chenopods are eaten (White 1994). In semi-arid woodlands, structural change in the vegetation community has occurred with an increase in the density of the shrub layer dominated by unpalatable shrubs such as Eremophila spp. , Dodonaea spp., Acacia spp. and a widespread loss of perennial grasses (Hobbs and Hopkins 1990).

The demise of Australia's mammals in the semi-arid and arid zones is also a result of European impacts. According to Morton (1990), of the 72 species of terrestrial mammals known from the arid zone, 11 are extinct, 5 have disappeared from the mainland and are confined to offshore islands and more than 15 have declined in range, persisting in semi-arid fringes but absent from the arid zone. Morton and Baynes (1985) have suggested that extinctions in the mammals preferentially occurred among species that had a specialised habitat or diet. Many arid zone animals remained relatively abundant through the 1920's and even later in the more remote areas but they rapidly -40-

declined after the introduction of pastoralism, feral animals and grazing stock. The decline and extinction of mammals is measured in decades and not centuries since European settlement (Recher and Lim 1990) attesting to a massive and sudden loss of species (Burbidge et al. 1988). An example of this is given in north western South Australia. Aboriginal people have confirmed that many species are no longer present in this region. From 43 species of indigenous mammals recorded, only 20 of these are considered extant. Within the last decade, 22 of the 103 terrestrial native species are presumed extinct in the state (Copley et al. 1989). More than one third of the terrestrial mammal species of the central desert region have vanished in the past 50 years, but Aboriginal people who lived in the Central deserts until recently maintain a knowledge of the mammals.

Morton (1990) proposes a conceptual model to account for the patterns of decline and extinction of mammals that is summarised below:

1) The environment of inland Australia was originally difficult for herbivores and omnivores because of infertile soils hence restricting the digestible production to small fertile patches. 2) The uncertain climate with droughts of irregular length contributed to this problem. As mammals were restricted to the scattered pockets of favourable habitat during droughts, this exposed them to high probabilities of local disappearances. Low (1984) referred to these as climatically limited species. The Lesser Stick-nest rat (Leporillus apicalis) is included in this category. Populations especially of small mammals may die in severely stressed refugia during prolonged droughts, and be replaced following a break in drought conditions by individuals that have survived, which are then able to radiate from the refugia during ephemeral plagues (Low 1984). 3) Grazing stock and rabbits upset the balance between the local disappearances and the reinvasions after a drought episode. The introduced herbivores maintained a higher population for a short time and altered the vegetational composition of habitats that the native species depended upon for refuge. As a result, the medium sized species are confined to much smaller patches of suitable habitat than larger species. Mammals are less mobile and it is more difficult to recolonise other areas after drought periods ease. 4) Degradation, introduced predators and altered fire regimes caused an increase in the probability of local disappearances and, several droughts later, resulted in extinctions.

Morton and Baynes (1985) suggest that the trophic level/position of the rodents made them particularly vulnerable to the impact of European settlement. This was a major reason for their extinctions in addition to other factors identified in Morton's (1990) model for mammal decline and extinctions. Species richness of rodents has declined to 44% of the Pre-European numbers except in the extreme south- west of the continent, -41-

and the species that survived tended to be small or inhabited rocky outcrops (Morton and Baynes 1985).

3.4 Semi-Arid Vegetation Communities: Sub-Continental Context

Factors that influence the composition and diversity of vegetation communities from the semi-arid zone have been described. The combination of these factors result in a spectrum of vegetation communities in the semi-arid environment: woodlands, shrublands, low shrublands, chenopod shrublands and tussock grasslands (Leigh and Noble 1969; Cunningham et al 1992). The flora consists mainly of sclerophyllous species with adaptations for survival in semi-arid environments that may include; phyllodes in the Acacia spp., viscid leaves in Dodonaea spp., tomentose leaves in Maireana spp., leaf abscission in Atriplex spp. and spines in Sclerolaena spp. (Stanley 1983). Features of each structural community will be outlined to illustrate the diversity of vegetation communities in the semi-arid zone. This knowledge sets the scene for understanding the spatial variability observed in the vegetation of the Flinders Ranges.

3.4.1 Tree and Shrub Species

There are no occurrences in South Australia of major natural forest or woodland formations classified as tall forest, closed forest or tall woodland. The reason for the lack of this vegetation type is climatic. Tall forests are not usually found in regions where mean annual rainfall is less than 1 000 mm. Only 1.15% of the state receives more than 600 mm of rainfall per year, which is regarded as a minimum for the growth of forests (Boomsma and Lewis 1980).

The genera Eucalyptus and Acacia are a feature of South Australian woodland vegetation. According to Boomsma and Lewis (1980) there are approximately 70 species and sub species of Eucalypts, 18 of which are tree species and the remainder are mallees. There are approximately 100 species of Acacia that range from understorey trees to arid shrubs. The next most common shrub genus in semi-arid vegetation communities is Eremophila (represented by 40 species) and understorey shrubs of Olearia (30 different species) and Hakea (20 species).

The initiation and development of buds on tree species are controlled by external factors such as good rain (Boomsma 1981). Some species of Eucalypts have a long flowering period, in contrast to other species such as Eucalyptus camaldulensis that have a short flowering period. Some Eucalypts have the ability to delay the opening of mature buds until a more favourable season arrives later in the year. However, most species would flower at least once over several seasons (Boomsma 1981). -42-

3.4.2 Chenopod Shrublands

Chenopods are perennial shrubs (Graetz and Wilson 1984) that most commonly include species from the Atriplex and Maireana genera (Oxley 1979; Stanley 1983; Beard 1984). Atriplex vesicarius is a short lived perennial shrub with a 10-25 year life span but only 12 months in periods of low rainfall (Williams 1979; Wilson 1990). It is opportunistic in flowering and seed set may occur several times a year, given sufficient rain (Williams 1979). The seeds have a longevity mechanism as they can lie dormant until favourable conditions develop. This species has the advantage of more rapid re- establishment by seed after a drought period breaks (Carrodus and Specht 1965). Maireana spp. are longer lived plants from 50 to 100 years, but the seeds survive less than one year in the field (Williams 1979; Wilson 1990). They are deep rooted, establish after heavy rains and flower prolifically in autumn/summer rains while seeding in spring and early summer (Williams 1979).

Chenopod shrublands have an understorey consisting of a sparse (canopy cover of <10%) ephemeral herbaceous layer (Beard 1984; Graetz and Wilson 1984; Wilson 1990). They cover 5-10% of the arid and semi-arid zones (Goodall 1979) with an annual rainfall of 150-200mm (Medlin 1993) and colonise xeric habitats, drainage lines, flood plains and depressions on saline and alkaline soils (Stanley 1983). This vegetation community can withstand drought periods of greater than 1 year and use rainfall at any time of the year (Walter 1971) but they flourish in the winter rainfall zone (Oxley 1979; Graetz and Wilson 1984). Ephemeral and perennial herbaceous species grow between the shrubs (Graetz and Wilson 1984; Wilson 1990) and are dominant in the community during dry periods but decrease in numbers during periods of high summer rainfall when grasses become more abundant (Medlin 1993). According to Stanley (1983), high shrub densities competitively exclude or suppress the growth of grasses and forbs (except in wet years).

3.4.3 Semi-Arid Woodlands

There are semi-arid and low semi-arid woodland vegetation formations, the former dominated by Eucalypts including Eucalyptus camaldulensis, E. microtheca and E. ochrophloia and the latter, Casuarina cristata, Heterodendrum oleifolium and Callitris columellaris (Stanley 1983). The Eucalypts are replaced by Acacia spp. when rainfall decreases (Walter 1971; Kuchel 1980). The understorey consists of an herbaceous layer containing a range of perennial grasses, ephemerals and perennial forbs (Wilson 1990).

According to Wilson (1990), the tree and shrub layer can be composed of approximately 20 different species at any one site and the total canopy cover may range -43-

between 5% and 30%. The shrub layer can vary from sparse to dense and in the adult stage, trees limit the growth of shrubs and together they limit the growth of perennial grasses (Harrington et al. 1984). Species in this layer may include Muehlenbeckia cunninghamii, Chenopodium spp. and a ground flora of perennial grasses such as Eragrostis setifolia and Paspalidium spp. (Stanley 1983).

3.4.4 Arid and Semi-Arid Low Woodlands

Northern communities in the semi-arid zone have a sparse cover of low shrubs and a ground flora dominated by grasses such as Aristida spp., Themeda australis and Chrysopogon fallax. The tall shrubs and trees in these woodlands include Casuarina cristata, Alectyron oleifolium and Heterodendrum oleifolium, a characteristic shrub layer of Eremophila sturtii, Senna spp. and Dodonaea spp. and a ground layer of Stipa spp., Sclerolaena spp. and Enneapogan avenaceus following summer rain (Stanley 1983).

3.4.5 Acacia Shrublands

These communities are often referred to as Mulga and range from low woodlands to tall shrublands and dense scrub (Stanley 1983). As environmental conditions become harsh, spaces between individual shrubs increase and the size of plants decrease (Boomsma and Lewis 1980). Mulga dominates 75% of shrublands and it is long lived to approximately 250 - 300 years. In the drier areas, seedling regeneration is episodic in response to rainfall (Stanley 1983). When summer and winter rainfall occurs, Acacia spp. can be replaced by Triodia spp. on sandy soils (Walter 1971), highlighting the influence of rainfall on the composition and structure of vegetation communities in semi-arid environments.

3.4.6 Grasslands

According to Medlin (1993) hummock/tussock grasslands found within the arid zone have an annual rainfall of 150-200 mm. Arid hummock grasslands are also known as spinifex grasslands and are dominated by species from genera such as Triodia, Plectrachne (Jacobs 1982; Medlin 1993) and Zygochloa (Medlin 1993). Trees, shrubs or any ground layer species are sparse or absent (Stanley 1983) but when present may include Acacia spp., Hakea spp., Solanum spp. and members of the Fabaceae (Griffin 1984). These communities form natural mosaics with shrublands throughout the Northern Flinders Ranges (Boomsma and Lewis 1980). They commonly occur on sand plains, dunefields and rocky ranges (AUSLIG 1990) and only exist as pure stands without a shrub over storey in the latter two environments (White 1994). Significant seed production occurs in years of extra rainfall (Jacobs 1982). -44-

3.5 Vegetation Communities in the Northern and Central Flinders Ranges: the Regional Context

Distribution of vegetation communities in the Flinders Ranges is controlled by geology and topography (Sprigg 1984, Greenwood et al 1989, Gell and Bickford 1996) that includes ridge tops, stony hills, valleys, gorges and outwash plains. These environments produce a mosaic of vegetation communities throughout the ranges (Fox 1991). Larger trees are found in sheltered areas and mallees in exposed sites with shrubs as understories. Exposed slopes have grasses and ephemeral species and fringing plains have saltbush shrublands (Swinbourne 1986). The principal vegetation formations in the region include dry sclerophyll woodlands (Callitris), riverine woodlands (Eucalyptus camaldulensis), mallee woodlands (Acacia), shrublands, chenopod shrublands and tussock grasslands (spinifex) (Beard 1981, Williams 1982). Vegetation communities are under the influence of shifts in the average effective rainfall and the season in which most of the rain falls (Gell and Bickford 1996).

Botanically, the ranges are an important region with species found there and nowhere else, for example Acacia araneosa (Swinbourne 1986). There are 85 plant species from the Flinders Ranges National Park that have been listed as having special conservation significance and 123 exotic species in the Flinders Ranges (Greenwood et al. 1989). The Northern Flinders Ranges (particularly 3 kilometres north of Arkaroola near Mount Painter) has a higher density of plant species of State and national significance compared to the central ranges (Greenwood et al. 1989).

Gell and Bickford (1996) have outlined vegetation communities from the Northern and Central Flinders Ranges adopted by the South Australia Geographical and Analysis Research Unit (GAR) and from previous mapping work by the South Australian Department of Planning and Environment. Features and indicator taxa of different communities are outlined in the following section.

3.5.1 Riverine and Dry Semi-Arid Woodlands

Woodland formations are found in creeks and gorges. Eucalyptus camaldulensis is the dominant species in River Red Gum woodlands that are confined to water courses (Greenwood et al. 1989, Beard 1981, Gell and Bickford 1996) and grow best on slightly acidic sandy or alluvial soils (Boomsma and Lewis 1980). Where drainage is impeded and there is semi-permanent water, species of theCyperaceae, sedges and Typha domingensis flourish (Gell and Bickford 1996). There can be a ground cover of herbs including Ptilotus obovatus, Sida petrophila, Abutilon leucopetalum and species of Asteraceae. Acacia victoriae and Bursaria spinosa shrubs have been recorded in the -45-

understorey of riverine woodands at Brachina Gorge. Melaleuca glomerata is more common in smaller creeks and gorges (Plate 3.1). Riverine woodlands are found along the banks and inside the channel of Arkaroola Creek, inside Mount Chambers Gorge and Brachina Gorge.

Low woodland communities have a variable composition of dominant species and understorey components. The dominant species include Callitris columellaris and Eucalyptus intertexta with perennial herbaceous plants and a greater abundance of grasses in the understorey in sheltered locations (Symon 1972, Boomsma and Lewis 1980, Sprigg 1984). On higher slopes, there can be an understorey of shrubs such as Bursaria spinosa, Dodonaea viscosa spp. angustissima, Senna artemisioides, Triodia irritans, Xanthorrhoea quadrangulata, Cassinia laevis and Olearia spp. Other understorey taxa of low woodlands may include Sida petrophila, Sida corrugata and grasses such as Danthonia spp. and Enneapogon avenaceus (Boomsma and Lewis 1980, Gell and Bickford 1996) (Plate 3.2).

3.5.2 Low Open Woodland

This community has dominant species found in Low Woodland formations as well as Casuarina cristata and Acacia aneura. Casuarina cristata occur as single trees, in clumps or groves (Boomsma and Lewis 1980). The understorey is more diverse including species such as Ptilotus obovatus, Sida calyxhymenia, Semicaluum spp., Eremophila freelingii, Pimelia microcephala, Enchylaena tomentosa and Rhagodia parabolica, Abutilon leucopetalum, Convolvulus erubescens, Euphorbia australis, Pterocaulon sphacelatum, Salsola kali, Sclerolaena divaricata, Trachymene glaucifolia and Trichodesma zeylanicum (Medlin 1993). Other indicator species include Acacia tetragonophylla, Solanum ellipticum and Acacia victoriae. Ground covers of grasses and exotic herbs such as Acetosa vesicarius and Echium plantagineum are features of this woodland community (Gell and Bickford 1996) (Plate 3.3).

3.5.3 Mallee Shrublands

Mallee communities have a patchy distribution throughout the Flinders Ranges and usually occur on rubbly shale valley sides and along established drainage lines (Boomsma and Lewis 1980; Sprigg 1984). The mallee zone is predominantly in the winter rainfall belt, receiving 250mm - 380mm of annual rainfall (White 1994). Eucalyptus intertexta and Eucalyptus socialis with Callitris columellaris, Casuarina cristata and Alectryon oleifolius are common species in the higher rainfall areas. A middle storey commonly includes Exocarpus aphyllus, Senna spp. and a range of saltbushes and bluebushes (White 1994; Gell and Bickford 1996). -46-

/ ...„. • *T-

0££! Plate 3.1: Riverine woodland community dominated by Eucalyptus camaldulensis in Brachina Creek at the western end of Brachina Gorge in the central Flinders Ranges.

Plate 3.2: Low woodland in the Arkaroola-Mount Painter Sanctuary dominated by Callitris columellaris. -47-

Acacia spp. occur in mallee communities with an understorey of shrubs including Eremophila spp., Dodonaea spp. and Senna artemisioides, and herbaceous plants such as Sida spp., Olearia spp. Ptilotus spp. and Chrysocephalem spp. that are prevalent after seasonal rain (Symon 1972; Sprigg 1984). Dodonaea spp. are noted for their ability to grow well on exposed rocky ridges and germination from seed is effective and continuous (Boomsma and Lewis 1980). The composition of the understorey varies according to the soil and habitat. For example, on sandy soils there may be Maireana spp., Enchylaena tomentosa, Senna spp. and a variety of grasses such as Enneapogon spp., Aristida spp. and Eragrostis spp. whereas in shallower soils there may be Senna spp. and Chenopodium spp. (Boomsma and Lewis 1980). Exotic species are also part of the understorey and include Acetosa vesicarius, Echium plantagineum and Solanum ellipticum. Mallee communities dominated by Acacia spp. are more widespread in the Arkaroola-Mount Painter Sanctuary as there is a greater abundance of rabbly shale valley walls (Sprigg 1984).

3.5.4 Tall Shrublands

These are dominated by Acacia victoriae and other co-dominant species including A. ligulata and A. tetragonophylla. A. victoriae has a widespread distribution in the Northern Flinders Ranges and will grow in a variety of habitats such as drainage lines, flood plains, slopes and valleys (Boomsma and Lewis 1980). The understorey is dominated by ephemeral herbs, Sida spp., Ptilotus obovatus. and Senna artemisioides are also widespread plants in tall shrubland communities growing in shallow soils on hill slopes (Boomsma and Lewis 1980; Kuchel 1980; Sprigg 1984) (Plate 3.4). Greenwood et al. (1989) also describe an open shrub community that is dominated by low shrubs, grasses and chenopods. The indicator species include A. victoriae, Dodonaea angustissima and Senna artemisioides with a ground cover of Sclerolaena spp. and Abutilon spp. Exotic species including Echium plantagineum and Acetosa vesicarius can be present.

3.5.5 Chenopod Shrublands

This community is found on the outwash slopes of the ranges in areas of lower rainfall and shallow soils (Gell and Bickford 1996) and poorly drained low salty depressions (Swinbourne 1986). Indicator species include Maireana pyrimidata, Atriplex vesicarius, Dissocarpus spp. and Chenopodium spp. The latter are often locally dominant in flood prone areas (White 1994). There are usually pasture plants in these communities that include Sclerolaena holitana and annual grasses such as Enneapogon polyphyllus, Stipa scalra and Themeda triandra (Sprigg 1984). Exotic ephemerals are present, but not as -48-

Plate 3.3: Low open woodland dominated by Acacia spp. on lower slopes and Callitris on higher ground near Arkaroola Creek in the Arkaroola-Mount Painter Sanctuary.

Plate 3.4: A mix of tall shrubland and open shrubland with Acacia and Eremophila spp., on a rocky slope in the Arkaroola-Mount Painter Sanctuary. -49-

*>-.""•" H$£»fcs : _-•»•*?

Plate 3.5: Chenopod shrubland community on the plain flanking the western side of the Flinders Ranges outside Brachina Gorge.

Plate 3.6: Upland tussock grassland community in the Arkaroola- Mount Painter Sanctuary -50-

dominant as in other vegetation communities. On higher ground, small stands of Acacia spp. and Casuarina cristata form part of this community (Swinbourne 1986) (Plate 3.5).

3.5.6 Tussock Grasslands

After the removal of grazing in the Flinders Ranges (approximately 1955), native tussock grasslands began to return (Bonython 1996), with taxa such as Enneapogon spp., Danthonia spp., Stipa spp. and Ptilotus obovatus. Dodonaea spp., Sida calyxhymenia and Solanum ellipticum are also indicator taxa for this community. High ridges are a feature of the northern Flinders Ranges and this topography is suitable for upland communities of tussock grasslands consisting most commonly of Triodia irritans and sparse Xanthorrhoea quadrangulata (Symon 1972, Kuchel 1980). Acacia victoriae may also exist as scattered shrubs in these communities in drier sites (Boomsma and Lewis 1980) (Plate 3.6).

3.6 Sensitivity of the Flinders Ranges: a refuge

A refuge can be considered in evolutionary or ecological terms. A region in which a suite of organisms can persist during a period when most of their original geographic range is uninhabitable due to climatic change is a refuge in evolutionary terms. There can be a high frequency of endemic species that are referred to as relicts (Brown and Gibson 1983). In ecological terms, a refuge implies a region where a suite of species persists for short periods of time when large parts of their preferred habitat become unavailable. This may be driven by climatic or ecological changes and these refugia may operate for less than one or up to a few generations (ERIN 1995).

According to a classification of refugia by ERIN (1995), the Northern Flinders Ranges was identified as a refuge on the basis of the number of endemic, relictual and other wise significant species present. Mountain ranges provide a mixture of refuges in evolutionary time, offering sheltered environments and geographically isolated habitats. They also provide water (from run off) and plant nutrients, producing resource rich areas that can function as refugia over ecological time frames. Greenwood et al. (1989) suggest that the ranges insulate vegetation from climatic change compared to the aridity on the flanking plains due to the availability of micro habitats. Gorges within mountain ranges are also refuge areas (ERIN 1995). They provide specialised microclimates for flora and fauna, allowing the survival of species that were more widespread in past wetter climates. This factor has implications for palaeoclimatic reconstructions for this region, as detection of vegetation change in pollen records from the Flinders Ranges is -51-

not only attributable to climatic change and vegetation within the refugia may be less affected by regional climatic change.

3.7 Regional Setting and Midden Study Sites: Implications for this Thesis

The Flinders Ranges is a region of diverse topography and habitats that have produced a mosaic of semi-arid vegetation communities in the landscape. In addition to this, it has been demonstrated that a variety of factors influence the structure and composition of vegetation communities from the semi-arid zone. Undertaking a palaeoecological study in a location such as this requires an awareness of this variability and its implications for individual midden records.

Middens are confined to certain environmental settings that are controlled by the geology and topography. Therefore, choice of study sites within the ranges and midden survey designs needed to take this into account. The variability of semi-arid vegetation communities was an important factor to consider in the design and location of sampling sites in both local and regional modern pollen studies. The spatial variability in pollen rain needed to be investigated on a local scale at midden sites in addition to understanding regional patterns in the pollen rain. Acknowledgement of the Flinders Ranges and its gorges as refugia, requires that midden sites are sampled from inside and outside these potentially sensitive local environments and from more than one locality in the ranges. This has implications for investigating the lag between vegetation response and climate change in these records and the reconstruction of palaeoclimates for this part of the semi-arid zone. The way this variability was accounted for in my sampling strategy is explained in the following chapter. -52-

Chapter 4: Methodologies: Strategies and Techniques for Midden Analysis

4.1 Introduction

There is a need for experimental studies of midden taphonomy whereby comparisons between midden contents, modern middens and local plant communities should be completed (Betancourt et al. 1990a). As established in Chapter 1, my work addresses two themes in midden analysis; methodological and taphonomic issues and the palaeoecological application of this type of record. Sampling strategies and techniques used in field work, and laboratory and data analyses are discussed in this chapter within the context of specific aims of this research.

A brief outline of the overall design of this midden research is followed by a description of the midden survey, techniques used to quantify and qualitatively describe vegetation communities at midden sites, and regional and local modern pollen studies. The next section outlines laboratory work involved in sub-sampling and processing middens and discusses how Accelerator Mass Spectrometry (AMS) radiocarbon dating was used to address methodological and taphonomic processes. The final section describes statistical analyses of midden records and modern pollen studies, designed to investigate temporal and spatial trends in the data.

4.2 Overview of Research Strategy

As explained in the previous chapter, any interpretation of palaeoecological records from middens must first account for the high spatial variability of the Flinders Ranges vegetation. I therefore decided to sample middens at three main locations within the Flinders Ranges to examine spatial and temporal variation via comparison of records from east-west and north-south transects.

The regional palaeoecological record will be derived from the analysis of pollen and macrofossil assemblages in middens. Temporal change will be investigated by analysis of a number of middens that span the Holocene and which are spread across sites in the Arkaroola-Mount Painter Sanctuary, at Mount Chambers Gorge and Brachina Gorge. Spatial comparisons can be addressed at each main study site as they are separated from each other by at least 100 kilometres.

The north-south and east-west comparisons are important for the interpretation of palaeoecological records as they will provide insight into the influence of the summer/winter rainfall boundary reflected in pollen and macrofossil records. -53-

An investigation of the regional and local modern pollen rain signals in relation to midden sites and present day vegetation cover has been derived from pollen traps along transects across the Flinders Ranges and from traps located at midden sites. These studies will give an insight into pollen recruitment and probable source areas of pollen taxa, which is important knowledge to successfully resolve methodological and taphonomic issues associated with midden analysis.

4.3 Arkaroola-Mount Painter Study Site

Arkaroola is the most northern study site in the Flinders Ranges (Figure 4.1). The Arkaroola-Mount Painter Sanctuary is 61 000 ha. in area and adjoins the Gammon Ranges National Park to the north east (Cook 1986b). North Well Creek, Radium Creek and Oppaminda Creek flow into Arkaroola Creek that drains north out of the Gammon Ranges, crossing low lying slatey hill terrain and back into deep gorges within the Arkaroola high granite country. It then swings eastward and the channel becomes sinuous with deep water filled rockholes, flowing out onto the plains and draining into Lake Frome (Sprigg 1984; Cook 1986b). There have been over 300 species of plants recorded at Arkaroola (Swinbourne 1986). High granitic ridges are a feature of the north and there is a distinctive upland community of tussock grasslands (Triodia irritans and Xanthorrhoea species) that colonise these areas (Symon 1972; Kuchel 1980). Callitris columellaris is also dominant in the Arkaroola region, growing in sheltered valleys and along rocky ridgelines (Sprigg 1984). Mallee communities are more widespread in the northern ranges on shale substrate in valleys.

4.4 Mount Chambers Gorge Study Site

Mount Chambers Gorge is located on the eastern flank of the Flinders Ranges situated between the Mulga-View Basin to the north west and the Frome Plains to the east (Medlin 1993), approximately 100km south of Arkaroola (Figure 4.1). On the southern side of the gorge entrance, is the bluff of Mount Chambers rising above broad river flats (Cook 1986b). Mount Chambers Creek twists through the gorge for approximately 18km to finally drain into Lake Frome (Cook 1986b). The gorge has been deeply incised by Mount Chambers Creek south of the East Chambers Range (Medlin 1993). Surrounding hills and the gorge are part of the western section of Wertaloona Station Pastoral Lease (Medlin 1993). Limestones are prominent in the stratigraphic sequence in the gorge, as capping on shales of high ridges or underlying valley floors and plains (Twidale 1980; Selby 1990). In some sections of the gorge, shale beds from the Tapley Hill Formation, form extensive scree slopes (Selby 1990). -54-

Lower rainfall and exposed stony ridges at Chambers Gorge have produced harsh environments for plant communities to colonise. North facing slopes (exposed to the sunlight for longer periods) tend to have a lower plant species diversity compared to the south facing slopes where lichens and ferns tend to colonise (Medlin 1993). Within Chambers Gorge, 54 vascular plant families are represented and 12% of plants are exotic species. The most common plant families are Chenopodiaceae, Asteraceae, Cruciferae, Poaceae, Mimosaceae, Malvaceae, Myoporaceae, Solanaceae and Zygophyllaceae. Along Chambers Creek and tributaries of Chambers Gorge there are open riverine woodlands. Along the margins of the main creek and better watered tributaries there are paper bark communities with Melaleuca glomerata and the less common northern paper bark Melaleuca dissitiflora (Medlin 1993). Casuarina cristata is an indicator species for low open woodland communities on slopes at the eastern and western end of Mount Chambers Range (Medlin 1993). Eucalyptus gillii mallee communities are located on the top of the East Chambers Range on dolomitic siltstones (Medlin 1993). Acacia shrubland communities occur on the lower slopes and hills surrounding Mount Chambers Creek and along banks of the main creek. Acacia salicina is a dominant species found in small scattered groups along the side of the main creek, particularly on the Main Bend and the northern slopes of Mount Chambers Range, west of the gorge. Outside the gorge there are chenopod shrublands with Atriplex spp. and Maireana spp. In the past few years, the Chenopodiaceae have declined and there has been an increase in introduced weeds on the Frome Plains (Medlin 1993).

4.5 Brachina Gorge Study Site

Brachina Gorge measures approximately 10km in length and was formed by the erosion and downcutting of Brachina Creek, draining the Aroona Valley and cutting through the Heysen and ABC Ranges (Cook 1986b; Mincham 1986) flowing westward towards Lake Torrens (Cock et al. in press). The morphology of the gorge is controlled by the basement rocks. The gorge at its widest diameter is in erodable siltstones and limestones (Proterozoic Wonoka and Bunyeroo Formations) and confined to a narrow gorge in resistant quartzose sandstones of the ABC Range (Wilpena Group) (Cock et al. in press). Brachina Gorge is approximately 70km west of Mount Chambers Gorge in the central Flinders Ranges (Figure 4.1).

Atriplex spp. and Maireana spp. colonise outwash plains to the east and west of Brachina Gorge. Inside the gorge, saltbush and shrubland communities colonise the rocky talus slopes. On lower slopes with more adequate soil development, low semi- arid woodlands dominated by Casuarina cristata, Alectryon oleifolium and Callitris columellaris (Stanley 1983), with an understorey of shrubs and a ground cover of -55-

Ltkt £fr»\ • Sticknest ral deposits f^72 Ranoes complex

Playa lake

O Settlement

—— Road J

Figure 4.1: Location of key stick-nest rat midden sites in the central and northern Flinders Ranges: Arkaroola, Mount Chambers Gorge and Brachina Gorge. -56-

ephemeral grasses and herbs are present. A patch of Eremophila platycarpum growing near a grove of Casuarina cristata trees occurs at the western end of Brachina Gorge. These species are commonly found in low open woodland formations as single or spreading trees over a low shrubby understorey (Boomsma and Lewis 1980). East of the stick-nest rat midden sites on steep slopes and ridges, there is upland tussock grassland vegetation and Callitris columellaris growing in sheltered positions on the slopes.

4.6 Midden Survey and Sampling

A systematic search for midden material was conducted over two field seasons in June 1994 and April 1995. Specific areas were chosen using local knowledge about known deposits. Other areas in the Flinders Ranges were searched based on the geology and occurrence of caves and overhangs in the landscape. Most successful search areas were rocky gorges and creek systems. Middens are poorly preserved in more open areas as they require suitably protected environments such as caves and overhangs to remain dry and fully preserved, and as has been found with Neotoma middens, preservation varies regionally due to the credibility of the substrate (Webb and Betancourt 1990).

Systematic surveys were completed along Brachina Creek in the gorge walls, the south western face of Mount Chambers and the walls of the gorge along Chambers Creek. Surveys from both of these sites were constrained along the main creek to ensure a representative sample of middens were located and collected. The Arkaroola-Mount Painter Sanctuary was more extensive geographically, and it was decided to target Arkaroola Creek and tributaries running west-east through the northern ranges for the midden survey. It was found that midden deposits were not as abundant in other areas of the Arkaroola-Mount Painter Sanctuary after conducting further surveys and sampling. The focus on Arkaroola Creek ensured a similar sampling strategy and research design to the other two study sites. The sampling strategy for the midden surveys ensured the spatial coverage of sites included an west-east and north-south dimension from gorges and creeks traversing the ranges and study sites extending from Arkaroola in the north to Mount Chambers and Brachina Gorges in the central Flinders Ranges.

The gorges and hills between Mount Chambers Gorge and Arkaroola would be a likely target for future midden surveys. The estimated totality of middens is largely unknown in the Flinders Ranges as this study is the first systematic survey. Leporillus spp. middens have been discovered by chance in association with owl pellet deposits at Brachina Gorge and Mount Chambers Gorge in work conducted by Mr. Graham Medlin. Fossil skeletal assemblages of mammals, lizards and birds are identified to investigate past fauna of the region (Medlin 1993). -57-

Middens collected from Arkaroola, Mount Chambers Gorge and Brachina Gorge were described, measured, sketched and photographed prior to sampling (see Chapter 6 and Chapter 7). Research Permits (U23214-02, W23215-02 U23214-04, W23215-04, U23214-05, W23215-05) were issued by The South Australian Department of Environment and Natural Resources under the South Australian National Parks and Wildlife Act, 1972 for sampling inside and outside reserves during each field season. Permission was given for collection of plant specimens from midden sites to aid in the interpretation of sub-fossil material, flowering plants for the pollen reference collection and minimum amount of midden sample required. When sampling middens where there was apparent layering in the deposit, the entire section was recovered in situ. Other middens were removed in sections using a hammer and chisel. Depending upon the midden size, shape and degree of induration, portions were removed from along fracture planes (Betancourt et al. 1990b) and therefore the weight (g) of each sample for analysis varied (Appendix 1). Samples were sealed in plastic sample bags and assigned a midden code and site name.

4. 7 Field Sampling of Modern Vegetation Communities

The primary aim of the modern vegetation sampling was to describe vegetation at each midden site qualitatively and to quantify the foliage cover and density of shrubs and ground cover within the vicinity of the middens. Quantifying vegetation at midden sites was required to make comparisons with the modern pollen data and to aid in the comparison of fossil pollen and macrofossil assemblages extracted from stick-nest rat middens. Goldsmith et al. (1986) state that measure of vegetation cover has been shown to be a sensitive method to record changes in the vegetation. This was achieved by the following strategy:

(i) at each midden site, plant species within a 100 metre radius of the cave or overhang was collected, identified and recorded. This produced a species inventory for each midden site within the expected foraging range of Leporillus.

(ii) three fifty metre belt vegetation transects were completed parallel to the slope contours at midden sites. Where possible, and safe access allowed, the transects were ran above the cave, at the cave/overhang opening and downslope of the cave with approximately ten metres between each transect (Dial and Czaplewski 1990). The transect midpoint was located at the rock shelter/cave opening (Cole and Webb 1985) and extended 25 metres either side of this. All species were recorded along the length of the transect and inside a one metre belt either side of the tape (Plate 4.1). This provided a species list at varying altitudes for each midden site. In addition to the transects, foliage cover and density for species that intercepted the transect lines were measured -58-

(McDonald et al. 1990; Cole and Webb 1985). Recording all species intercepting the transect is an acceptable method to use in areas where the vegetation cover is sparse (Kent and Coker 1992).

4.8 Pollen Trap Studies

Pollen trap studies were designed to investigate regional and local patterns of modern pollen deposition in the Flinders Ranges. The only modern pollen study from this vicinity was conducted by Singh and Luly (1991) at Lake Frome on the eastern boundary of the Flinders Ranges. This environment received a large proportion of regional pollen in the modern signals. The Flinders Ranges environment was expected to yield modern pollen signals that demonstrate its habitat diversity and consequently more variable vegetation. It should be more sensitive to local variations in individual taxa.

Modified Oldfield pollen traps consisting of a funnel secured into the neck of a plastic bottle were used. The funnel was lined with filter paper and synthetic fibre and a fly screen cap fitted over the top (Plate 4.2). Traps were buried in the ground along west- east transects (Plate 4.3), rocky slopes outside midden cave sites, and those located inside caves were anchored by packing stones around the base of the plastic bottle (Plate 4.4). Traps were set up for two six-month seasons: June 1995 to November 1995, and December 1995 to May 1996. Samples of modern pollen from winter and summer seasons served to minimise local bias (Kodela 1990) and detect uneven plant distributions that may result from different proportions of summer versus winter taxa.

The regional pollen signal along west/east transects across the Ranges was examined at Arkaroola, Mount Chambers Gorge and Brachina Gorge. I chose to place pollen traps at 5 km intervals along each of these transects to determine the pollen signal from different vegetation communities on the plains, flood plains, waterholes, gorges and cliffed outcrops.

The percentage ground cover of plant species within 4x4 metre quadrats surrounding the traps located in shrublands was recorded. In locations dominated by trees or vegetation in excess of 2 metres, the number of trees, species, height and trunk diameter were recorded within a 10 metre radius of the trap. According to Mueller-Dombois and Ellenberg (1974), this quadrat size is commonly used for woody undergrowth/shrub vegetation. -59-

If-* ^3 -Iff *** •BjJJw ^^

IKSsBuMk l?!

Plate 4.1: Vegetation transect ran parallel to the slope contour outside a midden cave site in the Arkaroola-Mount Painter Sanctuary. Species inventories and the foliage cover of species intercepting the transect tape were recorded.

naffim£3fcM

Plate 4.2: An example of a pollen trap used for both the west-east transect and midden cave pollen studies. -60-

Plate 4.3: Pollen trap located along the west-east transect across the Flinders Ranges in the Arkaroola-Mount Painter Sanctuary.

Plate 4.4: Pollen trap anchored inside a midden cave site in the Arkaroola-Mount Painter Sanctuary. -61-

I decided to set up pollen traps at each of the eighteen midden sites both inside and outside the caves/overhangs for the local modern pollen investigation. The deposition of modern pollen occurring inside and outside cave environments was monitored in order to investigate pollen taxa likely to be blown into the cave and incorporated into the midden, versus the pollen introduced as a result of midden construction and rat foraging activities. Determining pollen recruitment and source areas for the modern pollen will improve the integrity of palaeoecological interpretation of midden pollen assemblages.

Following each six month period in the field, funnels and contents were retrieved, bagged and bought back to the laboratory. Ninety three traps were set up over both seasons and 73 were recovered and processed. Table 5.1 and Table 5.13 in Chapter 5 provides specific details on recovered and lost traps. The most successful recovery was at Arkaroola and Mount Chambers Gorge. Trap losses were more variable at Brachina Gorge possibly due to the increased accessibility of sites to tourists.

4.9 Laboratory Methods

Midden processing and analysis in the laboratory was designed to investigate methodological and taphonomic issues. Fifty sub samples from the 18 Leporillus middens and 73 pollen traps were processed. Sub-sampling middens in the processing stage and AMS radiocarbon dating of components was undertaken to investigate the depositional history and timing of accumulation of these deposits.

4.9.1 Stick-nest Rat Midden Macrofossil Recovery

The processing of stick-nest rat middens followed established methods pioneered in Arizona USA. These were designed for the palaeoecological analysis of Neotoma spp. (packrat) middens. The aim of the processing was to separate, sort and identify macrofossils incorporated into the middens. According to Spaulding et al. (1990) and VanDevender (1990), to release the macrofossils from the amberat (urine) coating, the midden should be soaked in distilled water. The disaggregated residue can then be wet screened through a sieve, air dried, weighed and sorted.

To investigate the depositional history of middens and intra-midden variability, each midden sample was weighed (Appendix 1) and depending upon the size and structure of the midden, sub sampled further into top, middle and base sections. The majority of samples were in the order of 300 - 500 grams. Where relevant, location of sub samples is illustrated on the diagrams in Chapter 6 and 7. Some of the middens were difficult to sub sample due to their amorphous structure, while others were able to be sampled along fracture planes, similar to US techniques used with Neotoma middens (Betancourt et al 1990a). Other middens, typical of those at the Mount Chambers Gorge site, were -62-

fibrous and friable and only required dry sieving. Each indurated sample was placed in a container filled with distilled water and left to soak to dissolve the crystalline urine matrix (amberat). After the first 24 hours, as the midden began to soften, liquid was decanted and discarded to remove contamination of the outside by modern material and the midden was then soaked again in more water. This process of soak and decant was repeated for 5 to 10 days (depending upon the individual midden) until it was totally disaggregated. Large twigs and stone/rocks incorporated into the midden matrix were removed at this stage.

Liquid from the first soaking of middens was decanted and discarded. According to Mehringer and Wigand (1990), this step in the procedure minimises the potential of contamination by modern pollen. Issues of contamination were more difficult to monitor with the younger fibrous midden deposits that only required dry sieving. Extra care in field sampling was undertaken to minimise this factor. That is, the outer rind of the midden was removed prior to bagging and labelling the midden deposit. Aliquots of fine sediment and decanted supernatant left from each of the midden screenings were placed into 50 ml centrifuge tubes and washed in distilled water. Pollen was extracted via standard pollen techniques as outlined by Kershaw (1980) and Moore et al (1991) (Appendix 2).

The residue from each midden sample was then sieved through 2 mm and 500 (im sieves, to ensure macrofossils and small macroscopic material were recovered. The residues from the 2 mm and 500 |d.m fractions were then air dried, weighed (Appendix 1) and placed in a tray to sort and identify macrofossils. A sample of the <500 ^im fraction was kept in solution for pollen concentration.

A magilamp was used to view the macrofossils and they were separated from the grass matrix of the midden using tweezers and a brash. The fine fraction was scanned for macrofossils using a VMT binocular dissecting microscope (4X magnification). The macrofossils were divided into categories; leaf fragments and entire leaves, unidentifiable leaf fragments, seeds, fruits, twigs, insect remains, bone, hair, teeth and faecal pellets. They were placed in labelled glass phials ready for identification.

4.9.2 Identification of Macrofossils

Leaf fragments and seeds extracted from the middens were identified by staff at the Adelaide State Herbarium, South Australia and seeds from middens were identified by botanists at the New South Wales Agriculture Seed Laboratory in Sydney. Animal remains including bone and faecal pellets were identified by Mr Graham Medlin, Research Associate at the South Australian Museum, Adelaide and Professor Michael -63-

Tyler from the University of Adelaide. Insect remains are currently being identified to develop another avenue of palaeoecological reconstructions using stick-nest rat middens.

Leaf/fruit macrofossils were commonly identifiable to genus level but only in some cases to species due to poor quality preservation of the fossils. This is in contrast to macrofossil assemblages from Neotoma middens that are more numerous and well preserved (see studies in Betancourt et al. 1990b). There was more success with the identification of seeds as an extensive reference collection available from the New South Wales Agriculture Seed Laboratory was used to match the fossils with reference specimens. The bone material included teeth, bones and a large proportion of fragmentary remains. In most instances, bone could be identified to genus but there was difficulty in gaining further taxonomic precision. Teeth were the most successful fossils to be identified as they are directly comparable to reference collections held in the mammal section at the South Australian Museum. Some of the bone was too fragmentary for any identification and this was a result of some of this material being derived from regurgitated owl pellets that had been incorporated into the midden deposits (Medlin pers. comm. 1996). The majority of samples could not be ascribed to any particular species but were found to have similarities to the rodent group. Typical sizes of different rodent groups were determined from existing skeletal material in the collections and the fossil material was matched as closely as possible to one of these groups (Medlin pers. comm 1996).

After sorting, the macrofossils from each category were weighed and reported as a percentage of the dry midden weight prior to any processing. Counts of well preserved macrofossils (complete specimens) were also reported when weights could not be determined (ie. less than O.Olg).

Conversion of macrofossil data to a relative abundance scale was completed by dividing macrofossil weights by weight of the midden sample prior to soaking and converting into a percentage. The relative percentages were then expressed on a 5 point relative abundance scale where 1= rare (<5%), 2= occasional (5-10%), 3= common (11-50%), 4= very common (51-79%) and 5= abundant (>80%) (O'Rourke and Mead 1985) and used in the calculation of macrofossil :pollen index values. The scale used for converting macrofossil weights into a relative scale by Anderson and Van Devender (1991) was not applicable to stick-nest rat material, as calculations were based on a midden sample of approximately 800 grams. The majority of Leporillus midden samples were 500 grams or less, prior to soaking. -64-

4.9.3 Pollen Recovery from Traps

Synthetic fibre and filter paper that lined the funnels were washed thoroughly in distilled water and 10% KOH solution and centrifuged. The residue was then placed in polythene centrifuge bottles and KOH digestion was completed. The standard acetolysis and dehydration procedures were then completed and trap sample pollen concentrates were mounted in silicon oil on microscope slides for counting and identification (Appendix 2).

4.6.4 Pollen Reference Material

Reference material for modern pollen was collected over the period of 1994-1995. The flowers from species were collected, identified and sealed in envelopes. This collection was done at each midden site, vegetation transect and quadrat survey. Plant identifications were confirmed by botanists at the Adelaide State Herbarium. The procedure for preparing pollen slides from collected flowers and/or buds of living plants only required acetolysis. Anthers were mashed up in a solution of 30% glacial acetic acid followed by acetolysis, an alcohol dehydration series and mounting in silicon oil (Appendix 2). Reference samples are held in the School of Geosciences Pollen Reference Collection at the University of Wollongong.

4.10 Pollen Counting and Identification

Pollen grain identifications were based upon the exine characteristics, location and number of apertures and the arrangement of grains. Most grains are monads (single grain) however some occur in groups of four (tetrads) or more (polyads). The diagnostic feature is the number and arrangement of pores and colpi. Colpi may be constricted or display smooth margins whereas pores may have an operculum within the pore that is similar in texture to the pore or display a different textured area from the rest of the grain that surrounds the pore (Boyd 1992). The main morphological features are present on the outer surface of the pollen grain. These range from spinules to exine sculpturing including ragulae and reticulum processes and these characteristics are also used in the identification process (Boyd 1992).

Boyd's (1992) regional pollen atlas provided an important resource for this study. Identification criteria are summarised in Appendix 3, which includes a description of pollen grains, an average size of the grains and likely major sources for this pollen from taxa recorded in the Flinders Ranges vegetation. Eucalyptus, Melaleuca and Leptospermum were identified in the Myrtaceae family based upon the shape and characteristic margins of the grains. Chenopodiaceae and Amaranthaceae were combined in the final pollen sum due to the similarity between the grains (spheroidal -65-

and polyporate grains). The Pollen Reference Collection held in the School of Geosciences at the University of Wollongong was also used to cross check pollen from midden samples and traps with reference samples. The unknown category included damaged/crushed pollen and grains that were unsuccessfully matched to specimens in the reference collection.

Fossil pollen was examined under an Olympus CH-2 light microscope with a x40 objective and xl5 eye pieces. Two hundred grains per slide from transects two fields of view apart were counted and identified. However in some midden samples where certain taxa were dominant, the total count was increased until 200 grains of other taxa were recorded. A minimum of 100 grains was counted for each trap sample. Raw and percentage counts of pollen were recorded by Counter 3.0 ran on a Macintosh platform. The pollen sum was the total pollen counted and included Unknown pollen grains (Appendix 4).

Pollen diagrams for midden samples and pollen traps were constructed using WellPlot 1.0b Programme (Zippi Software) run on a Macintosh platform and Cricket Graph. Pollen taxa were also grouped into categories of trees, other Myrtaceae, tall shrubs, shrubs, herbs, grasses, chenopods/amaranths and exotics (Appendix 5). Taxa were classified according to the dominant life growth form of the plant recorded in field observations and in consultation with species lists compiled at the midden sites and comprehensive publications on semi-arid floras (eg. Cunningham et al. 1992). It is acknowledged that some taxa span these chosen boundaries so the structural categories on pollen diagrams need to be interpreted with caution.

4.11 Accelerator Mass Spectrometry (AMS) Radiocarbon Dating of Macrofossils and Pollen

Van Devender et al. (1985) have outlined special applications in the ability to date very small samples of plant macrofossils from Neotoma middens with Accelerator Mass Spectrometry (AMS) radiocarbon dating. The technique provides direct dates on plants of biogeographical interest in order to verify the presence or absence of different vegetation communities during the Late Quaternary. Justification of the use of AMS dating is based on the need for obtaining ages for very small samples (less than 200 mg) and the level of precision required (± 50 years) for improved temporal resolution of midden deposits. It has been suggested by Spaulding et al. (1990) that radiocarbon dating cannot resolve some taphonomic issues because at best the age only brackets the time of deposition. Van Devender et al. (1985) suggest that contemporaneity of macrofossils is reasonable, if the sample was collected from a discrete stratigraphical unit, any outer weathering rind removed and assuming that the ancient packrats were -66-

not fossil collectors themselves. My research tests this assumption whereby specific midden components will be individually dated, to investigate the contemporaneity of different components in the midden records.

Accelerator Mass Spectrometry (AMS) radiocarbon dating technology was developed in the 1970s enabling the relative numbers of ^C, l^C and ^C atoms in milligram samples of carbon to be counted (Brown et al. 1992; Arnold 1995; Litherland and Beukens 1995). This technique provides a major increase in the sensitivity of radiocarbon dating as the actual atoms are counted and the required sample size is smaller. The precision of AMS dates is dependent on ^C counting statistics of the sample and standards. Small contributions to the overall error of a date are due to the uncertainty of ^C and ^C measurements and therefore the sample is measured many times to minimise errors (Litherland and Beukens 1995). Contamination of a sample can occur at the site or during chemical preparation in the laboratory. The preparation is divided into two stages; pretreatment where the extrinsic contamination is removed through chemical extraction and the residue is combusted to CO2 followed by the conversion of CO2 to graphite (Litherland and Beukens 1995). Terrestrial plant macrofossils use atmospheric CO2 and are reliable material for dating and it is this material that is available from midden deposits.

I used Accelerator Mass Spectrometry (AMS) radiocarbon dating in this research to (i) measure the time span over which large amberat deposits from the Flinders Ranges accumulated in order to provide well dated palaeoecological records from single sites, (ii) compare ages of different components of the midden (e.g. faecal pellets, plant macrofossils and pollen) at discrete layers in order to examine taphonomic processes

Middens from Brachina Gorge (BR1, BR7, BR3, BR4) and the Arkaroola-Mount Painter Sanctuary (HH1, RC3, ARK1) were sub-sampled along fracture planes to retrieve top, middle and base sections. Other middens were amorphous and a representative sample from each midden was excavated.

4.11.1 Pollen Concentration for AMS Radiocarbon Dating

It is possible to directly date pollen extracts so that there is a direct association between the obtained dates and the palaeoclimatic indicator being studied (Brown et al 1992). The major challenge in dating pollen concentrates is obtaining a sufficiently large sample of pure pollen (Regnell 1992). Long et al (1992) have stated that 102 - 103 pollen grains (depending upon the species) are required for an acceptable AMS ^C date. It is a selective sample preparation technique that isolates material of interest from the contaminants; pollen from the organic residues. Sample preparation techniques vary -67-

in some details but basically use a modified standard pollen analysis and chemical treatment combined with additional sieving of the sample. The methods may be considered a further extension from pretreatments normally used in palynological studies (Brown et al. 1989). The technique is designed to remove the inorganic and non pollen material in that it exploits the chemical inertness of fossil pollen and allows for the selection of the size fraction of pollen to be dated via the choice of different sieve apertures (Brown et al 1992). The important difference is that chemicals containing modern carbon are avoided, by bleaching less resistant organic material with NaOCl (Sodium Hypochlorite) instead of using the traditional acetolysis treatment (concentrated H2SO4 and acetic anhydride). Compared with the traditional acetolysis treatment, the NaOCl is efficient in deflocculating and oxidising the amorphous debris to allow the separation of pollen during the sieving steps (Brown et al. 1992). The pollen concentrates are composed of pollen grains and small amounts of organic and cellulose material (Brown et al. 1989).

Brown et al (1989) and Regnell (1992) outlined two variations in the methodology used for the extraction of pollen concentrates (Appendix 6). My investigation used a modified version of these procedures due to resources available and laboratory safety issues working with Hydrofluoric (HF) acid.

The pretreatment step of the procedure consisted of 5cm3 samples of midden amberat being boiled in 6% KOH for 20 minutes and sieved at 180|um. The less than 180|im fraction was treated with cold HF (in place of a hot HF treatment) for 24 hours and then in hot HC1 for 10 minutes. The pollen was then concentrated by bleaching the pretreated residues with 2-3% of NaOCl for 5 minutes. The fraction was then sieved at 88|im and bleached with NaOCl for 5 minutes. The fraction was then sieved at 44|im and bleached again with NaOCl and sieved at 20|xm, retaining the 20-44u,m fraction. This was submitted for AMS radiocarbon dating at Beta Analytic. The sieving was done with fine nylon screens as opposed to metal sieves as they have been shown to be more effective in concentrating pollen from fine grained sediments as metal screens have a tendency to clog (Cwynar et al. 1979).

4.12 Statistical Analyses

There are temporal and spatial trends apparent in both modern pollen data and the stick- nest rat midden records that need to be investigated. Tests were required for comparison of macrofossil and pollen records from middens, assessment of inter-midden and intra- midden variability in fossil records and spatial patterns in the modern pollen rain. -68-

Cluster Analysis Cluster analysis assumes no external classification scheme but attempts to display groups and patterns inherent in the data as it objectively defines groups (Grimm 1988). In short, this technique summarises large data sets and aids in the interpretation of patterns within the data (Gauch 1982). When applied to vegetation data, this technique can designate major groups that correspond to modern vegetation units (Anderson and Davis 1988). The result is a two dimensional representation of the relationships between samples as a dendrogram. The analysis begins by using the centred data matrix to calculate a matrix of correlation coefficients between pairs of samples. The correlation matrix is searched for the sample pair exhibiting the highest correlation and when identified, the two samples are linked together. Samples thus linked are considered to be a single pooled sample. The output from the cluster analysis has required no information about the samples other than the data. Therefore cluster analysis permits objective comparison of all samples in the study, without regard to stratigraphic or geographic positions (Adam 1974).

Wards Hierarchical cluster analysis was used for modern pollen data sets. Pollen taxa were grouped into structural classes (trees, tall shrubs, shrubs, herbs, chenopods/amaranths and grasses) for the analysis. Pollen traps from the west-east transect studies were analysed to determine similarities between traps from different locations and determine regional pollen signals that are characteristic of the Ranges and surrounding plains. The analysis was also completed using all pollen taxa (less unknowns, exotics and other Myrtaceae) to check if local pollen assemblages influence the cluster analysis results.

Significance Tests Chi-squared significance tests were used for modern pollen data collected from traps located inside and outside caves at midden sites. These tests were used to investigate the differences in pollen signals inside as opposed to outside the cave environment. Tests were also completed on 1995 and 1996 traps from each midden site to determine whether there were significant differences between summer and winter trap pollen rain.

Similarity Indices and Macrofossil:Pollen Indices Sorenson Similarity Index (SI) values were calculated to quantify differences between taxa recorded in the modern pollen rain and taxa in the modern day vegetation cover at cave sites and trap sites along the west-east transect study. The application of this technique is suited to the presence/absence data that is used. The index is a ratio of the number of shared taxa in the modern pollen assemblage to the total number of taxa in both modern pollen rain and vegetation. -69-

Pollemmacrofossil indices were calculated for all middens to compare pollen and macrofossil assemblages. The data were scaled so that quantitative (pollen percentage) and qualitative (macrofossil) data sources could be compared (King and Van Devender 1977). According to Thompson (1985), the index should vary from -5 (taxa represented at levels greater that 50% in the pollen data and lacking plant macrofossils) to +5 (taxa are abundant in plant macrofossils and absent from the pollen). This calculation was described in Chapter 2. In addition to this index, the presence and absence of taxa from midden pollen, macrofossils and modern pollen were compared to an inventory of species present at the midden sites and taxa recorded from the vegetation surveys. This assisted with the interpretation of pollemmacrofossil indices and also provided insight into the relationship between pollen signals (fossil or modern) and the current vegetation cover,

4.13 Conclusion

This chapter has outlined the methodologies used in the field and laboratory followed by statistical analyses of midden records and modern pollen data. The choice and sampling of stick-nest rat midden deposits has allowed for detailed analysis of fossil material from Arkaroola, Mount Chambers Gorge and Brachina Gorge in addition to the opportunity to investigate the regional palaeoecological records by combining data from all stick-nest rat midden sites.

Vegetation surveys and modem pollen studies were designed to understand recruitment processes and source areas of modern pollen. This has implications for unravelling methodological and taphonomic processes that influence interpretations of midden palaeoecological records. The application of AMS radiocarbon dating in conjunction with a detailed sampling strategy of middens contributes to the resolution of methodological issues. And finally, statistical techniques will be used to investigate spatial and temporal trends in modem pollen and fossil records.

The following chapters will present results from the field and laboratory work. Vegetation transect and survey data will be described in the modern pollen trap studies (Chapter 5) in order to provide a modern analogue for the interpretation of palaeoecological records that are presented in Chapter 6 and Chapter 7. -70-

Chapter 5: Investigation of Modern Pollen Rain in the Flinders Ranges

5.1 Introduction

Understanding the modem pollen rain in semi-arid environments is an essential foundation for the interpretation of fossil pollen evidence. Issues considered in this chapter include the nature of modem pollen dispersal and deposition in a semi-arid environment. Results from regional and local studies, monitored from June 1995 to May 1996 in the central and northern Flinders Ranges, are presented. Cluster analysis and chi- square significance tests are used to analyse spatial and temporal variation in the modem pollen rain. Sorenson Similarity Indices were calculated to measure the degree of similarity between the presence of plant taxa recorded in pollen rain and the vegetation cover.

5.2 Understanding Modern Pollen Rain

The Australian semi-arid zone is a unique physical environment in terms of rainfall unpredictability, variety of topographic environments and the composition and structure of vegetation communities. In the Flinders Ranges, variable geology and topographic settings lead to high spatial variability in the vegetation as explored in the themes of Chapter 3. In addition to this, perennial, ephemeral and annual plant species and their associated responses to semi-arid rainfall patterns affect pollen production and consequently, the modem pollen spectra. Vegetation communities in the Flinders Ranges are under the influence of shifts in the average effective rainfall and seasons in which most of the rain falls (Gell and Bickford 1996).

5.2.1 Pollen Production, Dispersal and Deposition

Pollen grains are resilient and produced in large quantities (Martin 1973; Faegri and Iversen 1989), are widely dispersed and recovered from significantly small samples (Martin 1973), their relative frequencies are comparable (Faegri and Iversen 1989; Moore etal 1991) and production is more or less continuous (Head 1989). Since not all plants produce the same amount of pollen and not all pollen is preserved equally well there is not a one to one correlation between the representation of a species in the vegetation and the pollen record ( Webb et al. 1981; Faegri and Iversen 1989; Boyd 1992; Williams et al. 1993).

Pollen is available for transport via wind, water and vectors and the attributes of the pollen grains influence dispersal patterns, leading to spatial variation of pollen types -71-

(Prentice 1988; Williams et al. 1993). Well represented taxa are usually wind pollinated (anemophilous) whereas silent taxa are most often insect pollinated (entomophilous) (Prentice 1988). There can also be a combination of the two methods of pollination.

5.2.2 Finding a Modern Analogue

Reasoning by analogy is fundamental to palaeoecology. Determining how closely analogous past biotic communities are to present day ones provides a measure of amount and rate of ecological change through time (Delcourt and Delcourt 1991). There is an assumption that the modem plant taxon and fossil pollen have equivalent physiological and ecological responses to the environment (Nix 1978; Rymer 1978). The reconstruction of vegetation has to take into account the processes involved in the production of fossil pollen from the original plant cover (Jacobson and Bradshaw 1981; Faegri and Iversen 1989; Moore et al. 1991). To account for this variability, modem pollen samples are collected from known communities to be used for the interpretation, providing a link between pollen and the vegetation (Kershaw 1979; Williams et al. 1993). Environmental factors such as topography, aspect and climate that influence pollen must be considered (Faegri and Iversen 1989).

5.2.3 Representativeness of Pollen Taxa

In pollen rain studies by Dodson (1982; 1983; 1986; 1988), taxa were referred to in terms of representativeness. Pollen occurring at higher values in samples than the surrounding vegetation or whose potential source was absent from the local vegetation was over represented. Under represented taxa were those whose pollen was only likely to be represented when sampling under a source plant or whose pollen was inconsistently represented or present near the sampling site but not recorded in the pollen sample. Taxa present in pollen samples and vegetation at comparable percentages were considered as well represented. According to Dodson (1988), pollen production and dispersal can be grouped as:

(i) Local pollen, representative of species growing within 10 metres of the study site and usually low pollen producers exhibiting poor dispersal. (ii) Extralocal pollen that travels and is deposited within 100 metres of the study site. (iii) Regional pollen that travels greater distances and is deposited in large numbers.

Local pollen dispersal accounts for over 70% of Australian taxa although the majority of Australian plants are anemophilous (Dodson 1988). Casuarinaceae, Eucalyptus, Cupressaceae and Poaceae are known to be regionally dispersed and generally well to over representative of source vegetation. Asteraceae, Apiaceae, Sapindaceae, Chenopodiaceae/Amaranthaceae and Cyperaceae are also documented as well to over -72-

representative of source vegetation (Dodson 1988). Some of the poorly dispersed taxa include Acacia, Convolvulaceae, Euphorbiaceae, Fabaceae, Goodeniaceae, Myoporaceae, Solanaceae and Rutaceae. Loranthaceae is insect pollinated (Boyd 1992) so not expected commonly in the pollen spectrum. Annual species may explain the absence of families such as Haloragaceae and Liliaceae in the pollen samples. Some families or genera that include exotic species such as Polygonaceae, Cruciferae, Boraginaceae, Plantaginaceae and Pinus, are regionally dispersed and well to over representative of source plants.

5.2.4 Sampling Sites and Interpretation of Pollen Assemblages

Abundance of plant species surrounding pollen collection sites can indicate possible sources for pollen taxa recorded in traps. The size, density and location of the study areas will affect the proportions of local, extra local and regional pollen types (Jacobson and Bradshaw 1981) and the spatial resolution that is attained in the pollen assemblages (Webb etal. 1978; Gaudreau etal. 1989).

Jacobson and Bradshaw's (1981) model of pollen dispersal and deposition predicts that as the tree canopy opens above sites of increased basin size, the proportion of extra local and regional pollen will increase due to increased importance of pollen transport by wind and rainfall from the atmosphere. Therefore the range of taxa in the traps may not directly indicate a local vegetation type but represent different species at varying distances from the sample site. According to Martin (1973), the pollen in surface sediments from "small" catchment areas is usually dominated by species expected in the local surrounding vegetation.

Vegetational heterogeneity and differential pollen transport and preservation will continue to limit achievable precision in reconstruction of vegetation (Webb et al. 1978; Parson and Prentice 1981). Many different patterns of species abundance can yield the same flux densities of pollen types. This implies that interpretation of single pollen diagrams can be ambiguous. To counter this problem, a network of pollen diagrams distributed throughout a region should be used. Averaging pollen assemblages over several sites implies spatial smoothing which filters out local vegetation variation and overrides differences in source areas of different pollen types (Prentice 1988).

A limitation of pollen studies in arid areas is that pollen preservation at sites is often atypical of the broader region (Dodson and Wright 1989). Despite the diversity of plants within communities, most pollen samples are composed of predominantly few taxa which are usually the well known dominants. Within arid environments there is, relatively, a limited number of pollen grains produced by the local vegetation, and large proportions of regional taxa can mask the signal from locally occurring taxa (Dodson 1989; Horowitz -73-

1992). In cave deposits, pollen spectra varies in different parts of the cave (Davis 1990) as a result of the connection between the direction of the cave opening, prevalent wind directions and the pattern of vegetation distributions in the region (Horowitz 1992)

These factors have been addressed in the sampling technique utilised here. Traps were located in a variety of vegetation communities along west-east transects to determine regional pollen signals in the Flinders Ranges. Other traps were located at midden sites to determine local pollen rain and the effect of topography and aspect on the pollen spectra. It is then possible to examine how the regional and local signals compare to determine whether broad scale vegetation patterns are being reflected in the modem pollen.

It is often stated that proportions of pollen types found in a sample do not reflect the real composition of the vegetation because the relative abundance of pollen types is determined by pollen production and dispersal. This study aims to determine the representativeness of pollen taxa in a semi-arid zone environment by comparing signals to surveyed vegetation at trap sites and comment on the consistency of the representation of taxa across the study sites. This is similar in concept to Dodson's work, comparing the representativeness of a taxa or taxon recorded in traps to the presence/absence of this species in the vegetation. Luly (1990) concluded from a modem pollen study from semi-arid south eastern Australia that mallee heathlands, Callitris woodlands, chenopod shrublands and riverine woodlands could be distinguished from the modem pollen rain. However, no relationship could be discerned between pollen production and rainfall. Nevertheless this study is unique in providing modem pollen studies from rocky environments in the semi-arid zone. In addition, there is an expectation of extra variability induced by the stick-nest rat context and the importance of working out pollen sources and representation of taxa in this very particular context. Comparing modem pollen and fossil pollen assemblages from middens is allowing for the fact that midden pollen consists of pollen signatures from natural processes in addition to extra variability from Leporillus spp. foraging and nesting activities.

This modem pollen study will also make a contribution to existing knowledge of the characteristics of modem pollen rain at midden sites, and the relationship to pollen spectra in middens. Anderson and Van Devender (1991) investigated modem pollen rain from moss polsters and soil scrapes at packrat midden sites in the Waterman Mountains Arizona, to assess the relationship of pollen to vegetation. Modem pollen presented a biased view of the vegetation where trees and shrub taxa were under-represented, sub- shrubs strongly over-represented, perennial grasses moderately to under-represented and annual species poorly represented. Scott and Bousman (1990) compared midden pollen spectra with pollen from alluvial sediments and modern surface samples from Blydefontein Basin in the Karoo shrubland of South Africa and concluded that hyrax -74-

midden pollen reflected the local vegetation. Markgraf et al (1997) showed soil and moss polster samples in caves recorded high levels of tree pollen that compared well with midden fossil pollen. In mountainous settings, prevailing winds can carry pollen upslope from lower elevation communities (Thompson and Kautz 1983) that also affects modem pollen spectra at midden sites.

5.3 Pollen Traps and Vegetation Data

Descriptions of the modem pollen rain from west-east transects and cave sites at Arkaroola, Mount Chambers Gorge and Brachina Gorge are presented. There was a six month sampling period for each trap: 1995 from Winter-Summer and 1996 from Summer-Winter. Table 5.1 summarises the location and recovery rates of traps for the duration of the study for each transect location.

Vegetation was surveyed at trap sites during a 1995 field trip, using two methods along each west-east transect across the Ranges. The percentage ground cover of species from quadrat data was measured at trap locations. For the traps located inside and outside caves/overhangs, vegetation lists and foliage covers of species were recorded along transects. At locations where the vegetation was dominated by shrub and or tree species (unsuitable for quadrat survey), the diameter at breast height, mean crown width and crown gap were measured (McDonald et al. 1990). These observations are referred to in the text and on schematic diagrams illustrating vegetation cover at trap sites. Laboratory methods for processing traps are referred to in section 4.9.3 and pollen identification, counting and graphing procedures are detailed in section 4.10 of Chapter 4. Calculation of Sorenson Similarity Index is referred to in section 4.12 of Chapter 4.

5.3.1 Arkaroola Pollen Trap Transect and Vegetation Data

Traps along the west-east transect were located at 5 kilometre intervals: west of Haematite Hill; Junction of North Well Creek and Haematite Hill, Nooldoonooldoona Water Hole, Junction of Radium Creek and Arkaroola Creek and Echo Camp. One trap was buried on the eastern side of Arkaroola Rd, approximately 5km south of the west-east transect (Figure 5.1a). Data collated on ground cover species and vegetation communities at each trap site are illustrated in Figure 5.3. Low open shrubland communities were present at the western end of the transect. Riverine woodland communities were present along the confines of Arkaroola Creek and tall shrublands with chenopod and herbaceous understoreys were present to the south of the transect.

Arkaroola Transect 1995 Cupressaceae was represented along the transect at levels between 4% and 12%. Eucalyptus was present at levels of less than 10% (Figure 5.2). Asteraceae was most -75-

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Figure 5.1: Location of pollen trap sites along west-east transects across the Flinders Ranges in the

A. Arkaroola-Mount Painter Sanctuary B. Mount Chambers Gorge C. Brachina Gorge -77-

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Pollen Taxa Trap 1/95 Trap 10/95 Trap 11/95 Trap 23/95 Trap 1/96 Trap 10/96 Trap 14/96 Casuarinaceae ' * * * * Eucalyptus * Melaleuca * * * Myrtaceae A * Proteaceae * Caesalpinaceae * * Acacia * * * Sapindaceae * Myoporaceae * * Fabaceae * * Solanaceae * * * * Euphorbiaceae * * * Epacridaceae * * Asteraceae * Apiaceae * * Liliaceae * * Malvaceae * * * Convolvulaceae * * Haloragaceae * * * * Onagraceae * * * Lythraceae * * Cmciferae * * Cyperaceae * * Pinus spp. * Echium spp. * * * *

Table 5.2: Pollen taxa recorded at levels of less than 1% in traps from the west-east transect modern pollen study in the Arkaroola-Mount Painter Sanctuary. 79

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abundant at the western end of the transect, ranging from 36% to 56% in Traps 1 and 23 respectively. Chenopodiaceae/Amaranthaceae declined from west (21 %) to the east (1 %) and south (13%). Poaceae was represented at levels of 3-4% along the transect. Fabaceae, Sapindaceae and Apiaceae were present sporadically (Figure 5.2), and all other taxa with recorded levels of <1% are listed in Table 5.2. Trap 11 at Nooldoonooldoona Water Hole was the exception in this transect with 91% Cupressaceae, <1% Eucalyptus, and 1% of both Chenopodiaceae/Amaranthaceae and Poaceae (Figure 5.2).

Arkaroola Transect 1996 Cupressaceae declined from west (46%) to east (8%) whereas Eucalyptus levels were <5% at Trap 1 (west) increasing to 28% at Trap 14 (east). Asteraceae increased towards the middle of the transect, ranging from 6% at the western end to 21% at Trap 10, and dropped to 10% at the eastern end (Figure 5.2). The Chenopodiaceae/Amaranthaceae signal was similar to Asteraceae ranging from 6% at the western end to 21% at Trap 10 and back to 12% in the east. Poaceae was consistently represented along the western end of the transect at 12%-13% while levels were lower (<5%) at the eastern end. Herbaceous and shrub taxa recorded at levels <1%, are listed in Table 5.2.

There were higher levels of tree and herbaceous taxa in the 1995 trap series while Chenopodiaceae/Amaranthaceae and shrub taxa levels were similar in both 1995 and 1996 traps. Poaceae was more abundant in the 1996 pollen spectra.

Comparison of modern pollen and vegetation. At trap 1, there was approximately 40% ground cover of Dissocarpus paradoxus (Chenopodiaceae) (Figure 5.3) compared to pollen levels ranging from (6%-21%), suggesting this taxon may be variably represented when within 10 metre of the sampling point. Acacia was poorly represented in the pollen rain when plants were present in the vicinity of the trap, whereas Poaceae could be considered as more consistently represented. An additional twenty pollen taxa were recorded over the 1995 and 1996 periods that were not recorded in the vegetation (Table 5.3).

At trap 10, there was less ground cover of chenopods (Figure 5.3) however the levels of Chenopodiaceae/Amaranthaceae pollen were similar to the signal in trap 1. Variable levels of Asteraceae pollen may have originated from the site in addition to an extra-local source. Seventeen taxa were recorded in the traps that were not present in the quadrat (Table 5.3).

At Nooldoonooldoona Water Hole, pollen from canopy vegetation (Callitris columellaris, Melaleuca glomerata and Eucalyptus camaldulensis) masked contributions from other taxa at this site. Herbaceous species including Sida petrophila, Chrysocephalum spp. and -82-

others (Figure 5.3) were all <5% in ground cover and were either absent or present at levels <1% in the pollen sum.

At the eastern end of the transect (trap 14) there was a definite increase in the diversity of taxa in the pollen rain. There was evidence of locally occurring species either absent or at low levels in the pollen rain. Santalaceae (Exocarpos aphyllus) and Myoporaceae (Eremophila spp.) were absent in the modern pollen but present in the vegetation (Figure 5.3). Both taxa have poorly dispersed pollen (Table 5.9). Sixteen additional taxa were recorded in the pollen but not the vegetation (Table 5.3).

In trap 23, Goodeniaceae and Euphorbiaceae were recorded in the vegetation but not in the modern pollen. There was less than 5% ground cover of Solanum ellipticum and Solanum petrophilum (Figure 5.3) and <1% of Solanaceae in the pollen rain. Seventeen other taxa were in the trap but not the surrounding vegetation (Table 5.3).

Sorenson Similarity Index (SI) values were calculated for modern pollen samples and vegetation data collected along west-east transects to assess the degree of similarity between the modern pollen spectra and vegetation cover in the Flinders Ranges. The maximum possible value of SI is 1, where taxa in both the modern pollen rain and vegetation are identical although values above 0.9 are rare (Anderson and Van Devender 1991).

Table 5.4 shows index values for the 1995 west-east transect in the Arkaroola-Mount Painter Sanctuary. The degree of similarity between the modern pollen and vegetation is relatively low as evidenced by small SI values less than 0.5. There was a larger number of taxa represented in the pollen spectra in contrast to the vegetation cover adjacent to the traps.

Arkaroola Transect Pollen Traps PT1 5km west Haematite Hill PT10 Junction North Well Creek and Haematite Hill PT11 Nooldoonooldoona Water Hole PT14 Echo Creek Water Hole PT23 Paralana Springs Road * pollen trap not recovered

Table 5.4: Sorenson Similarity Index for comparison of taxa represented in the modern pollen rain and vegetation cover at trap sites along the Arkaroola-Mount Painter Sanctuary west-east transect for the sampling time of vegetation. -83-

5.3.2 Mount Chambers Gorge Pollen Trap Transect and Vegetation Data

Pollen traps were located along a transect beginning 5 kilometres west of Mount Chambers and finishing outside the eastern end of Chambers Gorge (Figure 5.1b).

Mount Chambers Gorge Transect 1995 Asteraceae declined from west (76%) to east (6%) along the transect whereas levels of Chenopodiaceae/Amaranthaceae ranged from 3%-15% outside the gorge and 18% inside the gorge (Figure 5.4). Poaceae was represented at levels between 5% and <1% and records of Cyperaceae were sporadic. Eucalyptus levels increased from west (<1%) to east (45%) and the same trend was observed for Melaleuca (Figure 5.4). Cupressaceae levels were <1% at the western end of the transect and increased to 7% in the east (Figure 5.4). Fabaceae signals were variable, ranging from <1% at the western end to 28% in the east. Signals from other herbaceous taxa were generally <1% (Table 5.5).

Mount Chambers Gorge Transect 1996 Eucalyptus increased from outside (5%-13%) to inside the gorge (35%-55%). Melaleuca showed the same pattern with 3% at the western end of the transect, increasing to between 6% and 9% inside the gorge. Cupressaceae was consistently represented with levels between 1% and 4% (Figure 5.4). Signals from shrub taxa were higher at the western end of the transect with Sapindaceae levels declining eastward and ranging from 9% to <1% and Fabaceae levels at 3%-6% outside the gorge compared to 1% inside (Figure 5.4). Asteraceae declined from west (32%) to east (6%) as did Chenopodiaceae/Amaranthaceae with levels at 31%-38% in the west and 15%-17% in the east. Cyperaceae was consistently represented with levels of 1% and Poaceae was more variable with levels between 1% and 8% (Figure 5.4). Other herbaceous taxa (with levels <1%) are recorded in Table 5.4.

Levels of tree taxa outside and inside Mount Chambers Gorge were similar during 1995 and 1996 while Asteraceae levels were higher in the 1996 traps and shrubs were more abundant in 1995. Signals from Poaceae and Chenopodiaceae/Amaranthaceae were similar during 1995 and 1996.

Comparison of pollen and vegetation Traps 9 and 10 at the western end of the transect were located in exposed sites. There was a sparse cover of Acacia shrubs and Poaceae in the vegetation (Figure 5.5), and these taxa were recorded in the modern pollen. An additional 23 taxa recorded in the trap but not in the vegetation (Table 5.6) must have come from a regional windblown source.

At the footslopes of Mount Chambers there was no Acacia or Proteaceae pollen in the trap although they were recorded in the surrounding vegetation (Figure 5.5). Both taxa are -84-

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Table 5.5: Pollen taxa recorded at levels of less than 1% in traps along the west-east transect modern pollen study at Mount Chambers Gorge. 86

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known to be poorly dispersed (Table 5.12). Another 14 taxa present in the modern pollen but not in the vegetation must have come from extra-local and regional sources (eg. Cupressaceae, Casuarinaceae, Eucalyptus) and ephemerals that might not be seen in any single year (eg. Onagraceae, Cruciferae) (Table 5.6).

Inside the gorge at trap site 11, Thymelaceae and Convolvulaceae were recorded in the vegetation at the site (Figure 5.5) and were poorly represented in the pollen (in 1996 only). There was 2% of Acacia pollen and Acacia victoriae was recorded within 10 metres of the trap (Figure 5.5). An additional 24 taxa were recorded in the pollen rain but not at the site (Table 5.6).

At the eastern end of the gorge (trap 12), all taxa present in the vegetation at the site (Figure 5.5) were also in the modern pollen. There was an additional 17 taxa in the pollen but not at the site (Table 5.6).

Outside the gorge (eastern end of the transect) there were 22 taxa represented in the modern pollen that were not recorded in the vegetation (Table 5.5). Pollen taxa that were present in the vegetation and modern pollen included Chenopodiaceae/Amaranthaceae and Convolvulaceae.

At Mount Chambers Gorge sites (Table 5.7), SI values were low at open trap sites emphasising a more diverse composition of taxa in the pollen rain with a dominance of regionally dispersed taxa. Index values at sites from inside Mount Chambers Gorge and at the eastern end where similar to each other (Table 5.7) but again there was a low degree of similarity between the pollen spectra and vegetation adjacent to the trap site.

Mount Chambers Gorge Pollen Traps 1995 PT9 5km west of Mount Chambers 0.08 PT10 Footslopes of Mount Chambers * PT11 Camp Ground Gorge 0.2 PT12 Chambers Gorge * 1 PT13 Eastern fence of Chambers Gorge 0.2 * pollen trap not recovered

Table 5.7: Sorenson Similarity Index for comparison of taxa represented in the modern pollen rain and vegetation cover at trap sites along the west-east transect at Mount Chambers Gorge. -89-

5.3.3 Brachina Gorge Pollen Trap Transect and Vegetation Data

Traps were located at 5 kilometre intervals along a transect beginning at the western boundary of the Flinders Ranges National Park and finishing east of the entrance to Brachina Gorge (Figure 5.1c).

Brachina Gorge Transect 1995 Two traps were recovered from the 1995 series (Table 5.1) and both were located at the eastern end of the transect. Cupressaceae (5%-9%), Eucalyptus (l%-5%), Acacia (3%- 5%), Asteraceae (3%-5%), Fabaceae (2%-4%) and Poaceae (2%-3%) were consistently represented at both trap sites. Chenopodiaceae/Amaranthaceae varied from 17% to 7% between the sites. Both trap sites also recorded high levels (compared to other transects) of exotic taxa ranging from 27% in trap 8 to 71% in trap 9 (Figure 5.6). The presence of Sapindaceae, Zygophyllaceae, Cruciferae and Cyperaceae were sporadic and other herbaceous taxa (Table 5.8) were present at levels <1%. Euphorbiaceae, Thymelaceae and Lythraceae were consistently low in the 1995 sampling period.

Brachina Gorge Transect 1996 Chenopodiaceae/Amaranthaceae levels declined from the western end of the transect, outside the gorge (61%), to inside the gorge (18%-20%), and increased again at the eastern end (31%) of the transect. Cupressaceae was consistently represented inside the gorge with levels between 3% and 6%, decreasing to <1% in trap 4 (Figure 5.6). Eucalyptus also displayed a similar trend with 16% at the western end of the transect increasing to levels between 23% and 28% further east into Brachina Gorge. Melaleuca was only recorded in this sampling period and Myrtaceae A was recorded at the western end of the transect at 2%-3% (Figure 5.6). Levels of Poaceae were consistently represented (2% and 4%) whereas the signal from Cyperaceae was more abundant at the eastern end of the gorge (2%-6%) compared to <1% at the western end (Figure 5.6). A variety of herbaceous taxa were present at levels of <5% and others at <1% (Table 5.8). iLevels of Chenopodiaceae/Amaranthaceae were higher in 1996 while tree taxa were higher in traps inside Brachina Gorge in 1996 compared to traps outside the gorge in 1995. There were similar levels of Asteraceae from both 1995 and 1996 traps while exotic taxa were less abundant in 1996 at the eastern end of the transect.

Comparison of modern pollen and vegetation. Approximately 60% of Chenopodiaceae/Amaranthaceae pollen (from a ground cover of 20%) and <1% Malvaceae pollen (from a ground cover of <1%) were recorded in the modern pollen, however Zygophyllaceae was only recorded in the vegetation outside the western end of the gorge at trap 4 (Figure 5.7). Zygophyllaceae is known to under -90-

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Pollen Traps Trap 8/95 Trap 9/95 Trap 4/96 Trap 3/96 Trap 2/96 Trap 8/96 Cupressaceae * Casuarinaceae * * Myrtaceae A * Myrtaceae B * Melaleuca * Leptospermum * * Proteaceae * Acacia * * Myoporaceae * * Sapindaceae * * Caesalpinaceae * Solanaceae * * * Euphorbiaceae * * Epacridaceae * Santalaceae * * Apiaceae * * Liliaceae * Convolvulaceae * Cruciferae * * Thymelaceae * * Malvaceae * * Zygophyllaceae * Haloragaceae * * * Onagraceae * Lythraceae * * * Cyperaceae * * Plantaginaceae * * Echium spp. * Unknown *

Table 5.8: Pollen taxa recorded at levels of less than 1% in traps from the west- east transect modern pollen study at Brachina Gorge. -92-

represent the source plant (Table 5.9). There were an additional 23 taxa represented in the pollen signal that were not present in the vegetation survey (Table 5.9).

Inside the gorge at trap 3, there was no Acacia pollen in the traps and Sapindaceae and Myoporaceae pollen were < 5%, however these taxa were recorded in the vegetation (Figure 5.7). Considering the crown cover of these shrubs, the pollen was under- represented. Chenopodiaceae/Amaranthaceae (20%), Asteraceae (<5%) and Thymelaceae (<5%) were present in the modern pollen rain and the vegetation, and there were an additional 17 taxa recorded in the pollen signal but absent at the site (Table 5.9).

Trap 2 was located in a patch of Eremophila deserti shrubs with a grove of Casuarina cristata trees within 10 metres of the trap (Figure 5.7). Myoporaceae and Casuarinaceae levels were < 2% in the pollen signal. An additional 19 taxa were recorded in the modern pollen but not at the site (Table 5.9).

Taxa recorded in both the modern pollen and vegetation at trap site 8 included Solanaceae, Eucalyptus and Chenopodiaceae/Amaranthaceae (Figure 5.7). There were 17 other taxa recorded in the pollen that were not present in the vegetation (Table 5.9). Higher levels of tree and Echium spp. pollen in 1995 may be a result of rainfall during winter and at the end of spring in 1995.

At trap 9, vegetation cover was sparse (Figure 5.7) and surrounding the site was chenopod shrubland and patches of mulga (Acacia aneura) within 20 metres. There was less than 10% Chenopodiaceae/ Amaranthaceae and <5% Acacia recorded in the modern pollen. However an additional 20 taxa recorded in the trap suggests an extra local/regional source. There was a large proportion of Echium in 1995 (consistent with trap 8) which may be a result of wet winter months enabling prolific flowering of this taxon (Table 5.9).

Brachina Gorge SI values (Table 5.10) were similar to those at Mount Chambers Gorge, given the low recovery of traps for the 1995 sampling period. There was more diverse representation of taxa in the modern pollen spectra irrespective of the vegetation cover adjacent to the trap sites. 93

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Brachina Gorge Pollen Traps 1995 PT4 Western end of the gorge * PT3 Brachina Gorge opposite BR4 * PT2 Brachina Gorge opposite BR1 * PT8 Brachina Gorge Gum Tree 0.18 PT9 Elatina/Nucaleena Formation 0.17 * pollen trap was not recovered

Table 5.10: Sorenson Similarity Index for comparison of taxa represented in the modern pollen rain and vegetation cover at trap sites along the west-east transect at Brachina Gorge.

5.4 Cluster Analysis of Transect Pollen Trap Study

A Wards Hierarchical cluster analysis (JMP statistical programme) was used to investigate spatial and temporal variability between modern pollen from Arkaroola, Mount Chambers Gorge and Brachina Gorge sites during the 1995 and 1996 sampling periods. The analysis was completed using all pollen taxa (less unknown grains, other Myrtaceae and exotics) from all traps.

Spatial Variability in the Modern Pollen The 1995 transects from Arkaroola, Mount Chambers Gorge and Brachina Gorge could not be distinguished from each other on the basis of all pollen taxa (Figure 5.8). There was a cluster of sites from similar vegetation communities at Arkaroola (A/Trap 1 and A/Trap 23), but other Arkaroola traps were more similar to sites at Mount Chambers and Brachina Gorge. Recovery of traps from the 1995 sampling period from Brachina Gorge was not as successful as 1996 and the analysis indicates that B/Trap 8 and B/Trap 9, both from the eastern end of Brachina Gorge, did not cluster. Traps from similar vegetation communities cluster together, irrespective of whether located in the central or northern ranges or from western or eastern positions along the transect.

Cluster analysis for the 1996 sampling period on all pollen taxa, grouped OTrap 11, 12 and C/Trap 13 (eastern end of Chambers Gorge) with B/Trap 4 (inside Brachina Gorge) (Figure 5.9) as a result of the dominance of Chenopodiaceae/Amaranthaceae pollen. It was not possible to distinguish differences between traps located in shrubland communities from each west-east transect, as sites at the western end of the Arkaroola transect (A/Trap 1, A/Trap 10) were grouped with sites at the western end of Mount Chambers Gorge (C/Trap 10, C/Trap 9) and a site at the eastern end of Brachina Gorge -96-

Clustering History Number of Clusters Distance Leader Joiner 8 5.5819170123 A/Trap 1 A/Trap 23 7 6.6048924414 A/Trap 10 B/Trap 8 6 6.6646354235 A/Trap 11 C/Trap 9 5 7.0654893726 A/Trap 11 B/Trap 9 4 7.6977800818 A/Trap 1 A/Trap 10 3 9.3409553547 A/Trap 1 C/Trap 11 2 9.6483505712 A/Trap 1 C/Trap 13 1 10.921499853 A/Trap 1 A/Trap 11 Dendrogram

A/Trap 1 A/Trap 23 A/Trap 10 B/Trap 8 C/Trap 11 C/Trap 13 A/Trap 11 C/Trap 9 B/Trap 9

Figure 5.8: Cluster analysis of modern pollen rain from all 1995 traps from west-east transects in the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge. -97-

Clustering History Number of Clusters Distance Leader Joiner 11 5.0701274151 A/Trap 10 B/Trap 8 10 5.6736060925 C/Trap 11 C/Trap 13 9 6.4759057164 C/Trap 11 C/Trap 12 8 7.5249760213 A/Trap 1 A/Trap 10 7 7.9569585858 A/Trap 1 C/Trap 10 6 9.3438582549 A/Trap 14 B/Trap 3 5 9.4578530923 C/Trap 11 B/Trap 4 4 9.4794538605 A/Trap 1 C/Trap 9 3 10.210361862 A/Trap 14 B/Trap 2 2 11.111512311 A/Trap 1 C/Trap 11 1 11.995510831 A/Trap 1 A/Trap 14 Dendrogram

A/Trap 1 A/Trap 10 B/Trap 8 C/Trap 10 C/Trap 9 C/Trap 11 C/Trap 13 C/Trap 12 B/Trap 4 A/Trap 14 B/Trap 3 B/Trap 2

Figure 5.9: Cluster analysis of modern pollen rain from all 1996 traps from west-east transects in the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and

Brachina Gorge. -98-

(B/Trap 8). This clustering was a result of similar signals of Asteraceae, Eucalyptus and Chenopodiaceae/Amaranthaceae in the modem pollen rain.

Temporal Variability in the Modern Pollen Cluster analysis was used to investigate differences between the 1995 and 1996 sampling periods for transects at Arkaroola, Mount Chambers Gorge and Brachina Gorge. All pollen taxa (less other Myrtaceae, exotics and unknowns) were used in the cluster analysis. At Arkaroola, the 1995 traps clustered separately from the 1996 series (Appendix 7). A/Trap 1:1995 and A/Trap 23:1995 were the most similar and A/Trap 1 and A/Trap 10 were recovered from both sampling periods and were in different clusters. At Mount Chambers Gorge it was possible to distinguish between the 1995 and 1996 sampling periods. C/Trap 12:1996 and C/Trap 13:1996 were the most similar to each other followed by C/Trap 11:1996 and C/Trap 9:1996 and C/Trap 10:1996 in a cluster from traps outside the gorge (Appendix 7). Traps from the 1995 sampling period are not similar to each other or the 1996 trap. Recovery of traps at Brachina Gorge was less successful and this hindered observations regarding temporal trends in pollen signals between 1995 and 1996. The 1996 traps were in a different cluster from the 1995 traps. B/Trap 8:1995 and B/Trap 8:1996 were not grouped in the same cluster (Appendix 7).

Chi-Square Significance tests were calculated on the composition of trees, tall shrubs, shrubs, herbs, grasses and chenopods for traps that were recovered in both 1995 and 1996 (ie. Arkaroola Trap 1 and 10, Mount Chambers Gorge Trap 9, 11, 13 and Brachina Gorge Trap 8). The 1995 and 1996 traps from individual sites were significantly different, indicating temporal variability (see Appendix 8 for Chi-square statistics and critical values). A cluster analysis was also completed for all traps (Arkaroola, Mount Chambers Gorge and Brachina Gorge) for both 1995 and 1996 years to investigate whether spatial or temporal variability was higher. The dendrogram indicates that generally 1995 traps cluster separately to the 1996 traps, irrespective of whether they are located at Arkaroola, Mount Chambers Gorge or Brachina Gorge (Figure 5.10). Two traps from Arkaroola 1996 transect (A/1:1996 and A/10:1996) clustered with the 1995 traps due to similar levels of chenopods and herbs. Two traps from Mount Chambers Gorge (C/l 1:1995 and C/13:1995) clustered with the 1996 traps based on more similar signals from trees and shrubs in each trap.

Differences between 1995 and 1996 trap sampling periods The cluster analysis has indicated that spatial variability in the modern pollen rain is large and it is not possible to clearly distinguish between Arkaroola, Mount Chambers Gorge and Brachina Gorge sites. However there is a tendency for traps located in similar vegetation communities (irrespective of the transect location) to cluster together, for -99-

Clustering History Number of Clusters Distance Leader Joiner 20 3.8571143324 C/13:1996 B/8:1996 19 4.6764915934 A/23:1995 B/9:1995 18 5.0332951406 C/9:1996 C/10:1996 17 5.79201613 A/1:1995 A/23:1995 16 5.9221617016 C/9:1996 C/13:1996 15 6.2658290296 A/1:1996 A/10:1996 14 6.7757749431 A/1:1995 A/10:1995 13 6.8265850346 B/2:1996 B/3:1996 12 7.4321804646 C/11:1996 C/12:1996 11 7.5430452992 A/1:1995 C/9:1995 10 7.5870981198 C/9:1996 B/4:1996 9 7.6166588955 A/14:1996 B/2:1996 8 8.3718095697 A/1:1995 B/8:1995 7 8.5437981879 A/11:1995 A/1:1996 6 8.8579331486 C/13:1995 0/11:1996 5 9.3125578687 A/14:1996 0/9:1996 4 9.8581252932 C/13:1995 A/14:1996 3 10.888291361 A/1:1995 A/11:1995 2 11.331337006 C/11:1995 0/13:1995 1 11.722033744 A/1:1995 C/11:1995 Dendrogram ~5 A/1:1995 A/23:1995 | B/9:1995 A/10:1995 C/9:1995 B/8:1995 A/11:1995 A/1:1996 A/10:1996 — C/11:1995 C/13:1995 C/11:1996 u C/12:1996 A/14:1996 B/2:1996 B/3:1996 C/9:1996 I C/10:1996 r^ C/13:1996 —1 B/8:1996 n B/4:1996 0

Figure 5.10: Cluster analysis of all 1995 and 1996 traps from west-east transects in the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge. -100-

example Arkaroola trap 11/1995 and Brachina Gorge trap 8/1995 in riverine woodland communities and Arkaroola trap 14/1996 and Brachina Gorge trap 3/1996 in shrubland communities. Temporal differences have been detected in the modern pollen over the 1995 and 1996 sampling periods at each study site. Discussion of possible explanations for this observation will be presented in the following section.

Monthly rainfall data for 1994-1996 from 6 regional rainfall stations was obtained from the South Australian Regional Office of the Australian Bureau of Meteorology. The corresponding transect locations for each rainfall station are listed (Table 5.11) and there are histograms of average monthly rainfall from 1994 to 1996 (encompassing the sampling periods of the pollen trap study) (Figure 5.11).

Station Number and Location Station Name Pollen Trap Locations 17099 - northern ranges Arkaroola Arkaroola 17052 - northern ranges Gammon Ranges (Wertaloona) Arkaroola 17054 - central ranges Gammon Ranges (Wirrealpa) Mount Chambers Gorge 17041 - central ranges Blinman Mount Chambers Gorge 19107 - central ranges Wilpena (Oraparrina) Brachina Gorge 19017 - central ranges Hawker Brachina Gorge

Table 5.11: Location and name of Australian Bureau of Meteorology regional rainfall stations in the northern and central Flinders Ranges

Monthly rainfall records indicate the wettest year in parts of the northern and central Flinders Ranges was in 1995 followed by 1996 and 1994. Stations under the influence of summer rainfall (17099 and 17052) recorded maximum falls from October to February in contrast to central stations under the influence of predominantly winter rainfall (17054, 17041, 19107 and 19017) with maximum rainfall occurring from May to July (Figure 5: 11).

The differences between 1995 and 1996 modern pollen can in part be explained by the rainfall records. Arkaroola, in 1995, shows high levels of Asteraceae and Chenopodiaceae/Amaranthaceae whereas in 1996 there were generally higher levels of tree taxa including Eucalyptus and Cupressaceae. Levels of Poaceae remained fairly consistent throughout the study. Rainfall records show a wet 1995 that was sustained into the summer months of 1996 (Figure 5.11) which would enable tree species to flower in the latter part of spring and therefore the signal could be picked up in the 1996 sampling period. The consistent record of Poaceae may be an indication of the summer rainfall influence over the Northern Flinders Ranges.

At Mount Chambers Gorge, there was overall less total rainfall compared to the other study sites, however we still see an increase in the level of tree pollen including -101-

Gammon Ranges (Wertaloona) 17052 Arkaroola 17099 125 -i 100-i

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Figure 5.11: Histograms of Australian Bureau of Meteorology monthly rainfall data (1994 to 1996) for six regional stations in the central and northern Flinders Ranges. -102-

Eucalyptus, Melaleuca and Cupressaceae and a drop in levels of Asteraceae in the 1996 sampling period, compared to the 1995 pollen rain. There was a relatively dry 1994 and a peak in the rainfall at the beginning of 1995, prior to the first trap sampling period. This was followed by an increase in monthly rainfall at the end of 1995 and into the early months of 1996 (Figure 5.11).

Rainfall stations near Brachina Gorge (Wilpena and Blinman) recorded high rainfall during 1995 and a lack of summer rain during the early months of 1996 (unlike the other stations). The majority of rain in 1996 was during the latter half of the year, characteristic of a predominantly winter rainfall regime. In the 1995 sampling period there were low levels of Poaceae and relatively high levels of Chenopodiaceae/Amaranthaceae and exotic taxa in the pollen rain. In 1996, the levels of tree taxa including Eucalyptus and Cupressaceae were higher and Chenopodiaceae/Amaranthaceae had also increased. The record of exotic taxa was most prominent in traps at the eastern end of the Brachina transect in the 1995 study. There was abundant rainfall during 1995, suitable for exotic taxa such as Echium spp. to flower and produce abundant pollen. This was prior to a marked decrease in exotic taxa levels during the 1996 sampling period, with relatively drier months up until the winter rainfall during the middle of the year.

5.4.1 Indicator Pollen Taxa

Based upon levels of taxa recorded in the modern pollen rain and comparison with vegetation cover at trap sites, Table 5.12 has been collated to indicate the representation of pollen taxa in the Flinders Ranges. This is similar in concept to Dodson's local/extra local/regional categories, referred to in section 5.2.3 and in this table. Representation of tree taxa including Eucalyptus and Melaleuca is variable between trap sites and levels are increased when species are present in the vegetation (Table 5.12). Cupressaceae is different in that it is over-represented in modem pollen irrespective of whether it is recorded in the vegetation. Levels of Casuarinaceae fluctuated between sampling periods at Arkaroola, and it was poorly represented in the pollen rain at Mount Chambers Gorge and Brachina Gorge. The diverse shrub taxa including Sapindaceae, Myoporaceae, Caesalpinaceae and Euphorbiaceae are generally under-represented in traps when compared to the common occurrence of these species in vegetation communities (Table 5.12), with contributions of <1% to the modern pollen spectra in open trap sites. The representation of Chenopodiaceae/Amaranthaceae also varies but this taxon occurs at consistently higher levels than other shrubs. Traps that were located in either chenopod shrublands or very exposed locations recorded higher levels of this taxon, indicating deposition from local and regional sources. There were lower levels recorded in woodland communities along the transects. Cyperaceae was poorly represented at trap -103-

Taxon Summary of pollen representation Acacia -poorly dispersed pollen -poorly represented in open trap sites along west-east transects when present in the vegetation -may be under-represented when vegetation is present at the site -only expected in the pollen rain if the source plant is within a few metres of the sampling site (Kodela 1990) Apiaceae -recorded in traps when not present in the vegetation at the site -higher levels recorded in cave trap sites -regional taxon in the modem pollen (Dodson 1988) -ranges from well to over-representative of source plants (Boyd 1992) Asteraceae -regional taxon (Dodson 1988) -well to over-representative of the source plant (Boyd 1992) -over-represented when herbaceous species dominate -well represented relative to abundance in vegetation at cave sites Boraginaceae -under-represented when herbaceous species dominate at cave sites -not recorded in open trap sites along west-east transects -are either annual or perennial species (Urban 1990) -well representative of source plant (Dodson and Myers 1986) Caesalpinaceae -under to well represented when present in the vegetation at cave trap sites -under-represented in open trap sites -perennial plants (Urban 1990) Casuarinaceae -under-represented in open trap sites -may be well represented at cave sites if present in the vegetation -regionally dispersed (Dodson 1988) anemophilous taxon (Green et al. 1988) -recorded as under to over-representing the source plant (Boyd 1992) Chenopodiaceae -tendency to be over-represented at trap sites in saltbush communities -over-represented in some cave sites when taxon is present locally -variable representation when tree or shrub genera close by trap sites -local and regional taxon (Dodson 1988) -well to over-representative of the source plant (Boyd 1992) and perennial species (Urban 1990) -104-

Convolvulaceae -under-represented when herbaceous species dominate in the vegetation at cave trap sites -poorly represented in open trap sites -plants are poorly represented by pollen (Dodson 1982) and perennial species (Urban 1990). Cruciferae -under-represented at cave sites when shrub genera dominate -recorded in open trap sites when not present in the immediate vegetation -insect pollinated annual taxon (Urban 1990) Cucurbitaceae -not recorded in the pollen rain at open trap sites or caves -perennial species (Urban 1990) Cupressaceae -over-represented when species is present in the vegetation at the site -over-represented at cave trap sites -regional (Dodson 1988) anemophilous taxon (Green at al 1988) -well to over represented (Dodson and Myers 1986) Cyperaceae -poorly represented at open trap sites when present in the vegetation -well to over-representative of the source plant (Dodson and Myers 1986) -poorly represented (Macphail 1979, Hope 1974 in Boyd 1992) Epacridaceae -under-represented in regional pollen rain at open trap sites -well represented when present in the local vegetation community that is dominated by herbaceous and shrub genera -some species are under-representative of source plants, while others are well represented (Boyd 1992) Eucalyptus -variable representation when vegetation is present in the immediate vicinity of open and cave trap sites - pollen is recorded when plants are absent in the vegetation -extra local or regional taxon (Dodson 1988) -variable representation of source plant (Boyd 1992) Euphorbiaceae -under representative of source plant when herbaceous taxa dominate in the vegetation -poorly represented in open trap sites Fabaceae -under represented in open trap sites -under representative of source plant (Boyd 1992) -annual and biennial species (Urban 1990) -105-

Geraniaceae -poorly represented at cave sites and regional signal and not recorded in the vegetation -poor to well representative of source plant (Boyd 1992) -short lived annual species (Urban 1990) Goodeniaceae -poorly represented at cave sites when present in the vegetation -under-represented in open trap sites -poorly dispersed (Dodson and Myers 1986) entomophilous perennial taxon (Green etal. 1988; Urban 1990) Haloragaceae -poorly represented in open trap sites -low levels recorded at cave sites in the absence of plants in the vegetation -source plants well to over-represented (Boyd 1992) Lamiaceae -poorly represented when shrubs are recorded in the vegetation at cave sites -poorly represented in open trap sites Leptospermum -low levels recorded in regional pollen signals in open trap sites -well to over -representative of source plant (Dodson and Myers 1986) Liliaceae -well represented at cave trap sites when herbaceous spp. dominate in local vegetation, otherwise levels are low -variable representation in open trap sites -varies in representation of source plant from under-represented (Dodson 1982) to well represented (Dodson and Myers 1986) -perennials that shoot annually (Urban 1990) Loranthaceae -poorly represented when present in the local vegetation under-represents to well represents source plants (Boyd 1992) -animal pollinated taxon (Urban 1990) Lythraceae -variable low levels recorded at cave trap sites -poorly represented in open trap sites Malvaceae -poorly represented at cave sites and open trap sites -under representative of source plant (Dodson 1983) -perennial taxon that germinates after rain (Urban 1990) Melaleuca -well represented at cave traps when recorded in the vegetation -insect pollinated taxon (Green et al 1988) -variable representation of source plants (Boyd 1992) -106-

Myoporaceae -poorly to under-represented at open sites when tree and herbaceous taxa are dominant -variable representation at cave sites when herbaceous spp. dominate -poorly dispersed (Dodson 1988) -over-representing the source plant (Dodson 1982 in Boyd 1992) Onagraceae -pollen under-represents source plants Other Myrtaceae -variable representation of source plants at cave sites likely from regional sources -poorly represented in open trap sites Pinus -poorly represented in open sites and cave traps Pittosporaceae -poorly represents source plants at cave trap sites -not represented in open trap sites -under or well representative of source plants (Dodson 1977b) -over-represents source plant (Dodson and Myers 1986) Plantaginaceae -poorly represented in open sites and cave traps -well to over-represents source plants (Dodson 1983 in Boyd 1992) ' Poaceae -variable representation in open sites and well represented at cave traps -regional taxon (Dodson 1988) -plants are well to over -represented by the pollen (Boyd 1992) Polygalaceae -not recorded in open traps or at cave sites -over-represents source plants (Dodson and Myers 1986) Polygonaceae -poorly represented in open sites and cave traps -wind pollinated (Boyd 1992) and well to over-representative of source plant (Dodson 1983) Proteaceae -poorly represented in cave traps when recorded in local vegetation -poorly represented in open trap sites -poorly represented (Hope 1974) to well represented (Macphail 1979) -animal pollinated (Urban 1990) Santalaceae -poorly represented when present in local vegetation at cave traps and open trap sites -under representative of source plant (Dodson 1977b) Sapindaceae -under-represented in open trap sites when herbaceous spp. and tree genera dominate -variable representation at cave trap sites -regional taxon (Dodson 1988) -well representative of source plants (Dodson 1982) -107-

Solanaceae -under-represented in cave trap sites when herbaceous taxa dominate and source plants are present in the local vegetation -poorly dispersed and under-represents source plants (Dodson 1977b in Boyd 1992) Stackhousiaceae -poorly represents source plants (Dodson and Myers 1986) Thymelaceae -poorly represented in open sites -under represented at cave trap sites -under-represents source plants (Dodson 1982) Typhaceae -poorly represented in open sites -well to over-represented by pollen (Dodson 1983 in Boyd 1992) Zygophyllaceae -poorly represented in open trap sites -under-represented at cave sites when chenopods and herbaceous taxa dominate the local vegetation -annual species that under-represents source plants (Dodson 1977b in Boyd 1992; Urban 1990)

Table 5.12: Representation of pollen taxon in the modem pollen rain from the northern and central Flinders Ranges, based on regional west-east transects and cave trap studies. Over-represented taxa refers to a strong representation of local and regional abundance and under-represented, a weak representation of local and regional abundance. -108-

sites while levels of Poaceae were variable at all sites along west-east transects at Arkaroola, Mount Chambers Gorge and Brachina Gorge. Herbaceous taxa were a variable component in the pollen rain. Most taxa were poorly represented at trap sites, with commonly low levels of taxa such as Convolvulaceae, Geraniaceae, Haloragaceae and Onagraceae (Table 5.12). There were a few exceptions including Asteraceae (over- represented when species were present in the local vegetation) and Apiaceae (recorded in traps irrespective of being present in the vegetation). There was a suite of taxa consistently represented at levels of <1% in the modem pollen. Solanaceae, Euphorbiaceae, Caesalpinaceae and Proteaceae were common taxa in the vegetation at trap sites but not abundant in the pollen rain. The diversity of taxa recorded at levels of <1% in the pollen rain was larger at sites from Mount Chambers Gorge, compared to Arkaroola and Brachina Gorge trap sites.

In summary, broad patterns of regional vegetation communities are being reflected in the modem pollen rain. Traps located outside gorges or in riverine woodland communities give distinct pollen signals from trap sites in shrubland or chenopod shrubland communities predominantly found on the rolling hills and plains flanking the ranges. Short term response of vegetation to different levels of available moisture (effective precipitation) is reflected in the modem pollen rain by increases in signals of Chenopodiaceae/Amaranthaceae and corresponding decrease in Asteraceae.

5.5 Pollen Trap Studies at the Midden Cave Sites

Results from the cave pollen studies at Arkaroola, Mount Chambers Gorge and Brachina Gorge are described in the context of vegetation recorded at the midden sites. The location of traps and recovery rate over the 1995 and 1996 sampling periods are summarised in Table 5.13. Details of the sampling strategy for the cave trap study are outlined in section 4.8 in Chapter 4. This section addresses what factors are determining the composition of modem pollen at this local scale, and how local and regional modem pollen spectra compare. The investigation of pollen rain inside and outside caves/overhangs provides an analogue for understanding pollen preservation in midden samples. Ultimately the question of whether pollen recruitment and preservation at midden locations is from local or regional sources must be addressed. This outcome will need to be taken into consideration when interpreting fossil pollen records.

5.5.1 Arkaroola Cave Traps and Vegetation Surveys

Haematite Hill Traps There were higher levels of Chenopodiaceae/Amaranthaceae, Cupressaceae and Eucalyptus outside the cave in trap 3/1995 (Figure 5.12). Levels of Poaceae remained -109-

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, Hakea ednieana , Dodonaea viscosa ssp onqustissima / Ptilotus obovatus s Triodia irritans ^TX -c f\ . Cal/itris columellaris Haematite Hill < < K , £remophla freelingii Midden / 5N& / • Dodonaea viscosa sip anqustistima . . Hakea ednieana xV v\ /.• DodonaeaCalhins columel/oriiviscosa ssp angustissima :?!?>->. / • Sida petrophtlo FOLIAGE COVER '••::'^*Ov / • Pfi lor us obovatus POLLEN TRAP • < IV. * "•':$>*>' • Triodia irritans • 1-5% yT ROCKY OUTCROP • 6 - 10% • II- 15 "A mSOI L ANO GRAVEL • 16-20% • > 20% ? 10

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i I I I I I lrl i i | i| i mi i i| i | I I | 1| L | I in 1995 in 1995 out 1995 out 1995 in 1996 in 1996 out 1996 out 1996 0 107c UJ

Figure 5.12: Pollen diagram of 1995 and 1996 traps (2 and 3) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour on Haematite Hill, Arkaroola-Mount Painter Sanctuary. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. -In­

consistent (inside and outside the cave) and there was a drop in shrub pollen (including Sapindaceae) outside. Apiaceae, Asteraceae and other herbaceous taxa were consistently represented in both traps for the 1995 sampling period (Figure 5.12).

Levels of Cupressaceae and Eucalyptus were higher outside the cave in 1996 (trap 3/1996). Asteraceae (29%) was higher outside and Chenopodiaceae/Amaranthaceae (10%-13%) was consistently represented. There was a higher level of Acacia (4%) and an increase in the diversity of herbaceous and exotic taxa outside the cave (Figure 5.12).

Trees were most abundant outside the cave for both sampling periods. Herbaceous taxa were the next most abundant group and levels were higher in traps inside the cave (Figure 5.12). Overall there was a more diverse assemblage of taxa in the 1996 traps with an increase in levels of exotics, chenopods/amaranths and grasses in the pollen spectra. The rainfall data (Arkaroola and Wertaloona) indicate a wet summer at the end of 1995 and continuing into 1996 with above average falls during March. Suitable conditions for flowering of spring-summer perennials and annuals at this time are reflected in the pollen assemblages.

Vegetation transects on Haematite Hill At the base of Haematite Hill there was a Callitris columellaris dominated woodland with an understorey of tall shrubs including Dodonaea viscosa, Acacia tetragonophylla, Acacia victoriae and less abundant Hakea ednieana and Senna artemisioides spp. sturtii. Rocky outcrops were colonised by scattered shrubs including Dodonaea viscosa spp. angustissima and Hakea ednieana. Downslope from the cave, foliage covers of shrubs and the diversity of understorey species increased (Plate 5.1). An herbaceous ground cover included Brachycome spp., Ptilotus obovatus, Sida petrophila and less abundant Solanum ellipticum. Cymbopogon ambiguus, Stipa spp. and Danthonia tenuior grasses were present in crevices on the rocky outcrops. Triodia irritans was growing at high elevations on rocky outcrops and further downslope on level ground. Foliage cover of species recorded along three transects at different elevations and parallel to the slope contour are summarised in Figure 5.12.

There were 13 taxa recorded in the pollen spectra inside the cave, and 14 taxa from outside that were absent in the vegetation (Table 5.15). Proteaceae was the only taxon present in the vegetation and not in the pollen, and Myoporaceae, Caesalpinaceae and Solanaceae pollen were only recorded in 1995. Pollen from these taxa are poorly dispersed and flowering usually occurs in spring-early summer. -113-

North Well Creek 1 Traps There were consistent levels of Asteraceae (50%-57%), Chenopodiaceae/Amaranthaceae (6%-10%), Cupressaceae (10%) and Apiaceae (8%-12%) both inside (trap 6/1995) and outside the cave (trap 7/1995). Levels of shrub pollen were higher outside the cave (4% Sapindaceae) and herbaceous taxa were consistently low (Figure 5.13).

In the 1996 sampling period, levels of Asteraceae were higher outside the cave (52%) and Chenopodiaceae/Amaranthaceae levels were consistent in both traps at 10% (Figure 5.13). There was a higher level of Eucalyptus recorded in trap 6 (inside the cave), however other tree taxa including Cupressaceae and Melaleuca were consistently represented (Figure 5.13). Shrub taxa including Acacia (4%) and Sapindaceae (3%) were more abundant inside the cave.

Herbaceous pollen was the most abundant signal both inside and outside NWCK1 cave followed by trees and chenopods/amaranths. The pollen assemblage from the 1996 sampling period was more diverse compared to 1995 (Figure 5.13).

Vegetation Transects at North Well Creek 1 Flowering Cassinia laevis and Eremophila freelingii were growing on rocky outcrops downslope and outside the cave. Other shrubs on the slope included Dodonaea viscosa spp. angustissima, Hakea ednieana and Senna artemisioides. Triodia irritans was the most abundant species at this site with a foliage cover of >20% on parts of the slope (Figure 5.13) growing with Sida petrophila and scattered Ptilotus obovatus, Solanum ellipticum and Solanum petrophilum. Marrubium vulgare and Bursaria spinosa were recorded on the mid slope sections in pockets of soil on the rocky outcrops. At 10 metres downslope from the cave there was a decrease in the foliage cover of Triodia irritans and Dodonaea viscosa spp. angustissima (Figure 5.13). Callitris columellaris trees were present on the lower section of slope, close to the creek and Cymbopogon ambiguus, Cassinia laevis and Melaleuca glomerata were growing in the creek bed (Plate 5.2)

There were 14 taxa recorded in the pollen inside the cave and 13 taxa outside that were not present in the vegetation at the site (Table 5.15). Lamiaceae and Pittosporaceae were present in the vegetation only and Malvaceae pollen was only recorded in 1995. These shrub and herb taxa have poorly dispersed pollen.

North Well Creek 2 Traps There was a high level of Cupressaceae (58%) recorded inside the cave in trap 4/1995 and weaker signals from Eucalyptus, Casuarinaceae and Melaleuca. Outside the cave, Cupressaceae levels were reduced (Figure 5.14). There were similar levels of Chenopodiaceae/Amaranthaceae and Asteraceae inside and outside the cave. Apiaceae -114-

Hakea ednieana Dodonaea viscosa ssp. anqusttssima Senna artemisiodies North Well Creek l.f| 3ida petrophita Midden Triodia irritans

Hakea ednieana Dodonaea viscosa Senna artemisiodies . Dodonata viscosa Sida petrophita . Cassinia laevis Trioaia irritans . Ptilotus obovatus • Triodia irritans Cumbopogon ambyuus FOLIAGE COVER T POLLEN TRAP • < IV. • I - 5% ROCKY OUTCROP • 6 - 10% • 11-15% SOIL AND GRAVEL • 16-20 % North Well Creek • > 20%

Trees Shrubs Herbs Grasses

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in 1995 out 1995 in 1996 out 1996 o 10% UJ

Figure 5.13: Pollen diagram of 1995 and 1996 traps (6 and 7) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at North Well Creek 1, Arkaroola-Mount Painter Sanctuary. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. -115-

Plate 5.1: Vegetation on the slopes of Haematite Hill 1 cave site.

Plate 5.2: Vegetation cover on the rocky slope outside North Well Creek 1 cave site. -116-

(49%) levels were higher in the pollen spectra outside (trap 5/1995) and there was a lower diversity of herbaceous taxa (Figure 5.14).

In 1996 Asteraceae (37%) and Chenopodiaceae/Amaranthaceae (16%) levels were higher in trap 5 (outside) whereas Eucalyptus (16%) was more abundant inside the cave. Melaleuca and Cupressaceae were consistently represented in both traps (Figure 5.14). There was a less diverse representation of shmb taxa outside the cave that included Sapindaceae and Solanaceae. Levels of Poaceae (4%) were higher in trap 5 whereas the representation of Cyperaceae was more consistent in both traps (Figure 5.14).

There was a strong signal from tree pollen inside the cave in 1995 whereas herbs, tall shrubs and shrubs were more abundant outside the cave. In 1996, herbs were the most abundant both inside and outside (Figure 5.14).

Vegetation transects at North Well Creek 2 A steep rocky slope rising to the top of a ridge was covered by Triodia irritans and scattered shrubs that included Eremophila freelingii, Dodonaea viscosa, Hakea ednieana and Senna artemisioides spp. artemisioides. The ground cover included sparse Sida petrophila, Goodenia vernicosa, Indigofera leucotricha and Solanum petrophilum. Cassinia laevis was present outside the cave entrance and further downslope amongst the Triodia irritan clumps (Plate 5.3). The foliage cover of Triodia irritans increased downslope with a reduction in the cover of Eremophila and Cassinia laevis (Figure 5.14). At the base of the slope, Ptilotus obovatus, scattered Abutilon leucopetalum and Solanum ellipticum were present.

An additional fifteen taxa were recorded in the pollen rain inside the cave, and not observed in the vegetation transects, and nine taxa in the pollen rain outside the cave that were not present in the vegetation (Table 5.15). Proteaceae, Goodeniaceae and Malvaceae, present in the vegetation, were not recorded in the modem pollen from either sampling period and are categorised as poorly dispersed pollen tending to under-represent source plants (Table 5.12).

Waterfall Traps Levels of Eucalyptus ranged from 15% to 41% over the sampling period and Cupressaceae and Melaleuca were more consistent. Chenopodiaceae/Amaranthaceae decreased in 1996 (9%) and Asteraceae remained consistent (Figure 5.15). The signals from Poaceae (14%) and Cyperaceae (5%) were more abundant in the 1995 sample.

Trees were the most abundant in the 1995 pollen spectra while there were similar contributions from herbs, grasses and chenopods/amaranths. In 1996 there was an -117-

"T^T S^-——- . hakea ednieana "V "* -r\ . £remophi/a free/mgri ~y^ . Qoodenia vernicosa TJ /, cassinia laevis -TT / . Sida petroohi/a North Well Creek 2^S^^ / . mddfa ir/i/ans Midden — —-^ ~^i ;V . £remophi/a free lman '•'•?\. . Dodonaea viscosa '•.'•'.^Sv . Cjoodenia vernicosa '•'•:-i\ /. Cassinia laevis \ "•ivN. / . 5ida petropMa TV / • Triod/a irritans

FOLIAGE COVER Y POLLEN TRAP T \ • < IV. "< ROCKY OUTCROP • 1 - 5% • 6 - 10V. • 11-15% P|]S0IL AND GRAVEL • 16-20 % 0 5 North Well Creek • > 20% motrei

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Figure 5.14: Pollen diagram of 1995 and 1996 traps (4 and 5) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at North Well Creek 2, Ark.aroola-Mount Painter Sanctuary. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. -118-

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Waterfall/ ^S~>- Midden' ^l-iisSv . Callitris columellaris £remophiia alternifolia i. Cassinia laevis / . Triodia irritans / . Cassinia laevis FOLIAGE COVER y POLLEN TRAP %? / • Triodia irritans < IV. -< ROCKY OUTCROP :.;:i • 1-5% _, • 6 - 10% QUJ SOIL AND GRAVEL -T>rr,:,; ,x • '"-ISV. g] TALUS 0 3 \ • 16-20 % • > 20% melrtt

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Figure 5.15: Pollen diagram of 1995 and 1996 traps (9) from outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Waterfall 1, Arkaroola-Mount Painter Sanctuary. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. -119-

increase in trees, herbs were similar to 1995 levels and grasses and chenopods/amaranths had decreased (Figure 5.15).

Vegetation transects at Waterfall site Vegetation surrounding the plunge pool at the base of the waterfall included Xanthorrhoea quadrangulata, Eucalyptus camaldulensis and Hakea ednieana. There were scattered Ptilotus obovatus, Sidapetrophila and Triodia irritans on the rocky substrate (Plate 5.4). Vegetation cover downstream from the plunge pool consisted of a ground cover of Triodia irritans and Cassinia laevis. There was a shrub layer of Eremophila alternifolia and Dodonaea viscosa spp. angustissima, and some Callitris columellaris (juvenile) saplings. Along the main channel the foliage cover of Triodia irritans increased (10%) and there was <1% of Cassinia laevis (Figure 5.15). Shrubs were also present and there was an increase in ground cover species such as Chrysocephalem semicaluum spp. semicaluum, Chrysocephalem semipapposum and Sida petrophila.

There were 16 taxa recorded in the pollen that were not present in the vegetation transects. Sapindaceae was present at the site and recorded in the pollen rain in 1996 only (Table 5.15).

Arkaroola Traps Levels of Cupressaceae ranged from 19% in trap 19/1995 (inside the cave) to 8% outside the cave (trap 20/1995) and Eucalyptus was more abundant inside the cave (8%). Levels of Asteraceae were higher outside and Apiaceae was consistently represented in both traps (Figure 5.16). Chenopodiaceae/Amaranthaceae ranged from 7% (trap 19/1995) to 34% (trap 20/1995) and Poaceae and Cyperaceae were consistently represented. There was a greater diversity of shrubs including Sapindaceae, Fabaceae, Solanaceae, Myoporaceae and Caesalpinaceae outside the cave (Figure 5.16).

There was a variable representation of Asteraceae in 1996 with 62% inside the cave (trap 19/1996) compared to 22% outside. Levels of Chenopodiaceae/Amaranthaceae (13%) were higher in trap 20 (outside the cave) as were Poaceae and Cyperaceae (Figure 5.16). Cupressaceae (41%) was abundant outside the cave with levels lower inside and other tree taxa were more consistently represented (Figure 5.16). There was a diverse assemblage of herbaceous taxa recorded in both traps whereas shrub taxa including Caesalpinaceae, Proteaceae, Myoporaceae, Fabaceae and Solanaceae were more prominent in trap 19 (Figure 5.16). Echium spp., Boraginaceae, Pinus spp. and Plantaginaceae were exotic taxa at the site.

In 1995, herbs were most abundant inside and outside the cave. Trees were the next most abundant inside whereas chenopods/amaranths were outside. Tall shrubs, shrubs and -120-

Plate 5.3: Triodia irritans (spinifex) covered slope outside North Well Creek 2 cave site.

Plate 5.4: Vegetation surrounding the plunge pool outside Waterfall 1 pi cave site.

'-M-'- ^ r;

Xanthorrhota evaofrnneju/ata • , Acacia iefraaonophyfla . PaoUtftaea wacasa ssp ongvatissima GoocUma vernicosa T" T\ • /« C»33ma tarn** 3 •f s /• CHrysoCMphofwn sesntca'uum r u ) / • Stafa pctrophifa / • Ptifotus ot>cratu3 Arkaroola l^«f Chenopodiuin spp T Midden ^T4\ • , Acacia tetraqoriophulla

t Acacia victoria* * Bremoph-la frtMl#lQfi i i, Qoodcrra verrucosa

;\ 1m Chrtfsoceptoatum msmtcatuum iCv.X / , PHtotuS Ohovatu* ":':;\ / • Callitris catumtltana '•K-lvN*,. 1 • Hak*a ednieana ''•"••'••'•v^s. • Acacia tetraaonophytla ''''•':'•:••w\. • Acacia victoria* "'""'••V^V^Nw^ /• £r*r*ophifa frselingii '"V.v^Sw / • Senna arttmisiodie* FOLIAGE COVER T POLLEN TRAP ^'v?^V / • CrOod€A&0 m 5 ^ >>*!«£$¥$' • 16-20% 0 • > 20% m«trti

Trees Shrubs Herbs

1 . J .J J> .# jf.jp t^Ss s •4? s/MP/j&ys *^S <•^& ^ ^ in 1995 out 1995 warn. • i in 1996 out 1996 •Mir mms i i Herbs Grasses

.'vi.p ///////cVf £

in 1995 I out 1995 in 1996 out 1996 o 10% UJ

Summary Diagram

in 1995 out 1995 in 1996 out 1996

Figure 5.16: Pollen diagram of 1995 and 1996 traps (19 and 20) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Arkaroola 1, ^karoola-Mount Painter Sanctuary. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. -122-

grasses were less than 10% of the pollen sum in both traps. In 1996, herbs were dominant inside and trees and herbs were most abundant outside. There was a drop in chenopods/amaranths in this sampling period (Figure 5.16).

Vegetation transects at Arkaroola/Devils Slide There was a low open shrubland community at this site, with an understorey of herbaceous taxa. Goodenia vernicosa was abundant on the slopes, with Sida petrophila, Abutilon leucopetalum, Solanum ellipticum, Chrysocephalem semicaluum and Cassinia laevis. Acacia tetragonophylla and Acacia victoriae were more abundant on sections of the lower slope (Figure 5.16). Eremophila freelingii was growing on the mid slope and other species present included Callitris columellaris, Exocarpos aphyllus and Sida petrophila. The foliage cover of Chrysocephalem spp. increased and there was less cover of Goodenia vernicosa and Ptilotus obovatus (Figure 5.16). Dodonaea viscosa spp. angustissima shrubs were growing outside the cave on the rocky outcrop with Xanthorrhoea quadrangulata (Figure 5.16). At the base of the slope there were Callitris columellaris trees and foliage covers of Acacia victoriae and Senna artemisioides spp. artemisioides increased (Plate 5.5). The ground cover was less diverse at this elevation.

An additional 12 taxa were recorded in the pollen inside the cave and 16 taxa outside the cave that were not recorded in vegetation transects. As Table 5.15 shows, Acacia, Proteaceae, Caesalpinaceae and Myoporaceae were recorded in only one year of the sampling period. This trend was observed at other cave sites.

Radium Creek 1 Traps Levels of Chenopodiaceae/Amaranthaceae ranged from 56% in trap 17/1995 (inside) to 11% in trap 18/1995 (outside the cave). Cupressaceae was consistently represented in both traps and levels of Eucalyptus ranged from <5% inside the cave to 7% outside (Figure 5.17). Asteraceae levels were higher outside the cave (24%) as were levels of Poaceae (14%) and Cyperaceae (8%). Herbaceous taxa including Malvaceae, Apiaceae, Convolvulaceae and Liliaceae were present in both traps (Figure 5.17).

The 1996 sampling period recorded an abundance of Other Myrtaceae pollen levels ranging from 22% inside (trap 17/1996) to 10% outside the cave (trap 18/1996). Eucalyptus levels were higher inside the cave (21%), Melaleuca was consistently represented at 5%-6% and Cupressaceae (13%) was higher outside the cave (Figure 5.17). Chenopodiaceae/Amaranthaceae levels ranged from 6% in trap 17 to 15% in trap 18, and Poaceae was more consistently represented between 5% and 7% (Figure 5.17). •123-

. Hakea ednieana / . £remophi/a fret/mg/i • Senna artemistodies . Sida petrophila . Solanum ellipticum , Triodia irritans T 1 T ( . £rvnophila fretlingii T T ) • Sida petrophila -tv*-<-

Radium Creek 1. Midden T/ T| T T T POLLEN TRAP T •< ROCKY OUTCROP • Citril/us lanatus 0 SOIL/ RIVER SAND

FOLIAGE COVER -^N*^^ Arkaroola • < IV. AT / > • 1-5% . • •^ >sw Creek • 6 - 10% • 11-15% • 16-20 % • > 20%

Trees Shrubs Herbs /// S/////* / rr Ss

Herbs Grasses

* .*,< v >v #y c^ ^ .1 in 1995 1 out 1995 in 1996 out 1996 0 10%

Summary Diagram

in 1995 out 1995 in 1996 out 1996

Figure 5.17: Pollen diagram of 1995 and 1996 traps (17 and 18) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Radium Creek 1, Arkaroola-Mount Painter Sanctuary. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1 % are listed in Table 5.14. -124-

Asteraceae (25%) was more abundant outside compared to inside (3%) and there were low levels of other herbaceous taxa (Figure 5.17/Table 5.14).

The composition of modern pollen inside and outside RC1 cave was variable. Inside the cave there were higher levels of trees and other Myrtaceae in 1996 and less abundant chenopods/amaranths (Figure 5.17). The signals from herbs remained similar but there were more shrub taxa present in 1996.

Vegetation transects at Radium Creek 1 Above the cave there was a sparse cover of Hakea ednieana, Eremophila freelingii and Senna artemisioides spp. sturtii and a ground cover of Sida petrophila, Solanum ellipticum and Triodia irritans. Solanum petrophilum, Abutilon leucopetalum, Galium spp. and Prostanthera striatiflora were present at this elevation. At the cave opening there was an increase in the foliage cover of Eremophila freelingii and Sida petrophila (Figure 5.17). The assemblage of ground cover was more diverse and included Ptilotus obovatus, Chrysocephalem semicaluum, Sida petrophila and Marrubium vulgare. Cymbopogon ambiguus was growing on the rocky slope and Stipa scalra spp. falcata, Brachycome ciliaris var. lyrifolia and Ixiolaena leptolepis were growing around the cave entrance. At the base of the rocky outcrop along Arkaroola Creek there was a riverine woodland community of Eucalyptus camaldulensis var. obtusa and Melaleuca glomerata (Plate 5.6). The Eucalypts had an average distance of 5 metres between the crowns and average crown diameter of 1.75 metres. An indeterminate species of the Cyperaceae were growing in areas of semi-permanent water. Citrillus lanatus (wild melon), an annual vine, and Pluchea dentex were present along sections of dry creek bed (Figure 5.17).

There were 12 taxa recorded in the pollen inside the cave and 3 outside that were not present in the vegetation (Table 5.15). Taxa recorded only in the vegetation included Proteaceae, Solanaceae and Curcubitaceae. Taxa only recorded in one season of the sampling period included Caesalpinaceae, Lamiaceae and Acacia (Table 5.15).

Radium Creek 2 Traps Levels of Cupressaceae (45%) and Eucalyptus (11%) were higher inside the cave (trap 15/1995) whereas Asteraceae (20%) and Chenopodiaceae/Amaranthaceae (22%) levels were more abundant outside the cave (trap 15/1995). Combined shrub taxa including Acacia, Sapindaceae, Myoporaceae, Solanaceae and Caesalpinaceae produced a variable and weak signal inside and outside the cave (Figure 5.18). Exotic taxa including Pinus spp. were recorded in trap 16/1995 (outside). -125-

•rt A

T/

Radium Creek 2 tg^ti T T Midden ^"^ TJ

T) FOLIAGE COVER • < IV. T[ • 1 - 5% • 6 - 10% • 11-15% T\ • 16-20 % • > 20% T| \ -# Melaleuca glomeraia 3fc SPECIES RECORDEO AS PRESENT (tea tail) T\ .Acetosa vesicarius Y POLLEN TRAP -A Ptilotus obovatus \ 1 Pluchea dentex < ROCKY OUTCROP 0 SAND AND GRAVEL metres Trees Shrubs Herbs

P J1 . «# * C G*^< <$ / •^ ^ SSJOWSSSS/cf~

in 1995 out 1995 in 1996 out 1996

Grasses

in 1995 out 1995 in 1996 out 1996 0 10% Summarry rDiagra m JT * faff cf'

Figure 5.18: Pollen diagram of 1995 and 1996 traps (15 and 16) from inside and outside the midden cave site and vegetation species outside the cave at Radium Creek 2, Arkaroola-Mount Painter Sanctuary. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. -126-

,*- " V.'' 4- • . *

-r ; - '4,-'' J* 1 Plate 5.5: Vegetation outside Arkaroola 1 cave site. Acacia victoriae, Eremophila spp.and a ground cover of herbs were common on the rocky slope.

Plate 5.6: Riverine woodland along Arkaroola Creek downslope from Radium Creek 1 cave site. -127-

Eucalyptus was not consistently represented with levels ranging from 37% inside (trap 15/1996) to 23% outside (trap 16/1996) whereas Cupressaceae (9%-12%) and Chenopodiaceae/Amaranthaceae (10%-13%) levels were similar in both traps (Figure 5.18). Levels of Poaceae, Cyperaceae and Asteraceae were more abundant outside (trap 16/1996). There were low levels of shrub and herbaceous pollen taxa in both traps (Figure 5.18).

There was an abundant signal from trees in the modern pollen both inside and outside the cave for 1995 and 1996. Herb and chenopods/amaranths signals were more abundant in 1995 whereas the signal from grasses was most abundant outside the cave in 1996 (Figure 5.18).

Vegetation at Radium Creek 2 The slope at RC2 midden site was an open rock face (Plate 5.7) and too steep for vegetation transects. Species within the vicinity of the midden included Melaleuca glomerata, Acetosa vesicarius, Ptilotus obovatus and Pluchea dentex.

There were an additional 18 taxa recorded in the pollen inside and 21 outside the cave, that were not recorded in the vegetation inventory (Table 5.15)

Radium Creek 3 Traps Eucalyptus was consistently represented in trap 12/1996 (inside the cave) and trap 13/1996 (outside the cave) with levels ranging from 31%-25% (Figure 5.19). Asteraceae (14%) was more abundant in trap 12/1996 and Chenopodiaceae/ Amaranthaceae levels (13%) were higher in trap 13/1996. There was a diverse representation of shrub taxa including 7% Myoporaceae, 5% both Sapindaceae and Fabaceae, 14% Caesalpinaceae and <5% Acacia and Solanaceae inside the cave. Recorded levels outside were less abundant (Figure 5.19).

Tree pollen was the most abundant inside and outside the cave followed by shrubs and herbs. Levels of grasses and chenopods/amaranths were higher outside the cave (Figure 5.19).

Vegetation transects at Radium Creek 3 Clumps of Triodia irritans were a dominant component of the vegetation community on this slope and the foliage cover increased on lower sections of the slope (Figure 5.19). Sida petrophila and Eremophila freelingii covers decreased and other shrubs on the lower section of slope included scattered Prostanthera striatiflora and Acacia confluens. There was a diverse ground cover of species including Olearia decurrens, Cymbopogon ambiguus, Ptilotus obovatus and Solanum ellipticum. Brachycome spp. and Eriachne -128-

Eremophila freelingii Senna artemmodiet Oleana decurrens Radium Creek 3 Sida petrophila Midden —' Solanum ellipticum Ptilotus obovatus Triodia irritans Curnbopooort amb/ouus

Acacia confluens £remophila freelingii Proatanthcra strtatiftora Chrysocephalurn semicoluum sida petrophila Solanum ellipticum Ptilotus obovatus Triodia irritans

FOLIAGE COVER T POLLEN TRAP <• IV. K ROCKY OUTCROP I- 5% [i]] SOIL AND GRAVEL 6 - 10% [2 BOULDERS ON TALUS SLOPE II- 15% 16-20% 4J. VEGETATION TRANSECT IS AT W THE SAME ELEVATION AS THE CAVE > 20%

Trees Shrubs Herbs Grasses

in 1996 out 1996

Summary Diagram $ & <& jf ^ X? &

*? K& *& till i i J_L_J ±JL J_l _L_L in 1996 wmmm i out 1996 0 10% uJ

Figure 5.19: Pollen diagram of 1995 and 1996 traps (12 and 13) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Radium Creek 3, Arkaroola-Mount Painter Sanctuary. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. -129-

mucronata were also recorded at this elevation (Plate 5.8). Cheilanthes austrotenuifolia was present at the cave opening in addition to Chrysocephalem semicaluum, Ptilotus obovatus and sparse Solanum ellipticum.

Seven taxa were recorded in the pollen rain but not in vegetation transects parallel to the slope. Proteaceae, Lamiaceae and Malvaceae, although present in the vegetation, were not recorded in the pollen rain (Table 5.15).

Oppaminda Track Traps Levels of Chenopodiaceae/Amaranthaceae ranged from 23% inside the cave (trap 21/1995) to 13% outside (trap 22/1995). There was a consistent representation of Eucalyptus (12%-13%) and Asteraceae (12%-17%). Cupressaceae (21%), Sapindaceae (8%) and Poaceae (13%) were more abundant in trap 22/1995 (outside the cave) (Figure 5.20). Herbaceous taxa were represented at levels of <5% inside the cave and included Onagraceae, Malvaceae and Cruciferae. A small pollen sum was recovered from trap 22/1995 (Appendix 4).

Trap 21/1996 (inside the cave) recorded 10% Cupressaceae, 9% Poaceae and Cyperaceae, 7% Chenopodiaceae/Amaranthaceae and 4% of both Eucalyptus and Fabaceae (Figure 5.20). Shrub taxa were represented by <5% each of Myoporaceae, Sapindaceae, Caesalpinaceae and Fabaceae. There was 3% Asteraceae, 2% Liliaceae and other herbaceous pollen were <1% (Figure 5.20 and Table 5.14).

In 1995, tree pollen was more abundant than any other group outside the cave whereas inside there were similar levels of tree, shrub, herb and chenopods/amaranths. Representation of grasses was stronger outside. In 1996 the trap recorded a signal from trees and similar levels of herbs, grasses and chenopods/amaranths (Figure 5.20).

Vegetation transects at Oppaminda Creek The Oppaminda Creek transect was run parallel to the cave entrance. There was a shrubland community with 1-5% foliage cover of Eremophila freelingii and Exocarp aphyllus (Figure 5.20). Other shrubs included Alectryon oleifolium, Hakea leucoptera with Convolvulus remotus in the foliage, Acacia victoriae and Senna artemisioides (Plate 5.9). Ground cover species included Ptilotus obovatus, Solanum ellipticum, Stipa spp., Convolvulus remotus and Marrubium vulgare. Cymbopogon ambiguus, Abutilon leucopetalum, Enchylaena tomentosa and a grove of Casuarina cristata were recorded within the vicinity of the cave

There were 13 taxa represented in the pollen inside the cave that were not recorded in the vegetation and 8 taxa from outside the cave (Table 5.15). •130-

Plate 5.7: Steep sparsely vegetated rocky slope outside Radium Creek 21 cave site. Melaleuca glomerata are present at the base of the slope.

Plate 5.8: Eremophila freelingii and Triodia irritans commonly found on the rocky slope outside Radium Creek 3 midden cave site. -131-

"^T-TT^

T] . Itlarrubiom vu/port. r . Eremophila Ihttlmgii r^\ . Casuarina cristata T_/ • Btocarpus ophyltus

Oppaminda Track ^^t^^^ • Ptilotus obovatus • iolanum ellipticum Midden T T* r • stipa spp- T TSs. • Convolvulus rtmotus

FOLIAGE COVER «. .„ T POLLEN TRAP • —> IT. • 1 - 5% J ROCKY OUTCROP •"•••" '•$?&''•" Oppaminda Creek • 6 - 10% n • 16-2011-15%% S 0 SOIL AND GRAVE5 L • '2°* m.

Trees Shrubs Herbs

* jf * & ^ .<•?• «s> JJW •? .JT J?

in 1995 out 1995 in 1996 FrrrrrFrrrrFFrn

Herbs Grasses

1 L_l l 1 1 1 J_l in 1995 1 • wsmm • I out 1995 >•• in 1996 1 • • f o 10% UJ Summary Diagram

^ «^ <3? # ^

I • i i i I—LI—1—IUI I II—I—'I I ' ' 1—' in 1995 •HH • imm mm* u wmm i out 1995 in 1996

Figure 5.20: Pollen diagram of 1995 and 1996 traps (21 and 22) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Oppaminda Track 1, Arkaroola-Mount Painter Sanctuary. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.14. -132-

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5.5.2 Mount Chambers Gorge Cave Traps and Vegetation Surveys

Mount Chambers 1 Traps The level of Chenopodiaceae/Amaranthaceae (86%) was abundant in trap 1/1995 (outside the cave) compared to other taxa. Shrubs included 2% of Sapindaceae and 1% of both Caesalpinaceae and Myoporaceae and tree taxa included <1% of Cupressaceae, Casuarinaceae and Eucalyptus (Figure 5.21 and Table 5.16). There was 3% of both Apiaceae and Asteraceae and lower levels of other herbaceous taxa (Figure 5.21).

Levels of Chenopodiaceae/Amaranthaceae were abundant both inside and outside the cave in 1996 with 84% in trap 1/1996 and 62% in trap 2/1996. Asteraceae was consistently represented with levels ranging from 4%-6% (Figure 5.21). Cyperaceae (2%) and Poaceae (2%) were more abundant outside, compared to the levels in trap 1/1996. Shrub taxa were more diverse in trap 1/1996 with contributions from Acacia, Myoporaceae, Caesalpinaceae, Sapindaceae, Fabaceae and Solanaceae (Figure 5.21).

Chenopods/amaranths were the most abundant category both inside and outside the cave. Outside in the 1996 sampling period, herbs were the next most abundant and there was an increase in the level of trees compared to 1995 levels (Figure 5.21). Late spring to autumn of 1996 was a wet period, which may explain an increase in the levels of herbs, grass and tree pollen outside the cave.

Mount Chambers 1 Vegetation Transects Vegetation was patchy on the talus slopes on the south western face of Mount Chambers (Plate 5.10). Outside the cave, the foliage cover of species included <5% of Acacia victoriae, 2% of Ptilotus obovatus and <1% of Sida petrophila, Chenopodium spp., Prostanthera striatiflora, and Atriplex spp.(Figure 5.21).

There were an additional 21 taxa recorded in the pollen but not the vegetation inside the cave and 12 taxa from outside (Table 5.17). Each taxon that was recorded in the foliage cover was also represented in the modern pollen. There was a more diverse assemblage of herbs, trees and grasses in the modern pollen. This site was exposed and conducive to regional and extra-local input of pollen blown in by wind.

Mount Chambers 2 Traps Chenopodiaceae/Amaranthaceae (41%) was more abundant in trap 4/1995 (inside the cave). There were higher levels of Cupressaceae (11%) inside the cave and Casuarinaceae was consistently represented (9%-6%). Less than 2% of Myoporaceae, Sapindaceae, Caesalpinaceae, Fabaceae and Solanaceae were in trap 4/1995. Asteraceae (11%-16%) -137-

Dodonaea microzuga Prosfanihera atriatifloro Mount Chambers I. Sidb pefrophila Middenv T Atriplex spp. Acaao victoriae TM Ptilotus obovatus Mount Chambers I (exposed) Midden

FOLIAGE C< T POLLEN TRAP . < iy. A. ROCKY OUTCROP . 1 -5% • 6 - 10% E) SOIL AND GRAVEL • II - 15% | GRAVEL AND BOULDER 16 • -20% ' TALUS SLOPE • > 20%

Trees Shrubs Herbs Grasses

£

mm I I I I I I I I I in 1995 in 1996 out 1996

Summary Diagram

Figure 5.21: Pollen diagram of 1995 and 1996 traps (1 and 2) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Mount Chambers 1, Mount Chambers Gorge. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.16. -138-

Plate 5.9: Eremophila freelingii shrubs growing outside the cave site at Oppaminda Creek.

Plate 5.10: Sparse vegetation cover outside MCI cave site on the south western face of Mount Chambers. -139-

was consistently represented and other herbaceous taxa such as Onagraceae (8%) was more abundant in trap 3/1995 (outside the cave) (Figure 5.22).

Chenopodiaceae/Amaranthaceae was consistently represented inside (trap 4/1996) and outside (trap 3/1996) the cave with levels between 43%-44%. Eucalyptus levels varied between 3%-7% (Figure 5.22) and Cupressaceae and Casuarinaceae were similar in both traps. Cyperaceae/Poaceae was consistently represented and Asteraceae ranged from 16% in trap 4/1996 to 35% in trap 3/1996. The signal from shrub taxa outside the cave was less diverse compared to trap 4/1996, with Acacia and Sapindaceae present (Figure 5.22).

In the 1995 sampling period, trees, herbs and chenopods/amaranths were more abundant than shrubs and grasses. In 1996 there was an increase in herbs and chenopods/amaranths (outside the cave) and a decrease in the signal from trees (Figure 5.22).

Mount Chambers 2 Vegetation Transects There was a moderate foliage cover of Olearia decurrens outside the cave entrance and less cover of Ptilotus obovatus and Poaceae spp. (Figure 5.22). Other species recorded along the transect included Acacia victoriae, Solanum ellipticum, Prostanthera striatiflora and Bursaria spinosa. Stipa spp. was growing out of the crevices surrounding the cave entrance. Ptilotus obovatus, Chrysocephalum semicaluum and Acacia victoriae were present ten metres downslope of the cave and other species recorded at this elevation included Casuarina cristata, Sida petrophila, Solanum petrophila and Prostanthera striatiflora (Plate 5.11).

There were an additional 16 taxa recorded in the pollen and not the vegetation inside the cave and 13 taxa outside. Trees and shrubs were growing in the vicinity of the cave that contributed to a more diverse vegetation community compared to site MCI. Lamiaceae and Malvaceae were only recorded inside the cave in 1995 and Pittosporaceae was only recorded in the vegetation (Table 5.17).

Chambers Gorge 1 Traps In pollen trap 5/1995 (downslope from Chambers Gorge 1 midden site) there was 21% Chenopodiaceae/Amaranthaceae, 17% of both Eucalyptus and Poaceae and 10% Cyperaceae. Shrub taxa included 7% of both Myoporaceae and Fabaceae and lower levels of Acacia and Caesalpinaceae (Figure 5.23). Herbaceous taxa were represented by Asteraceae (3%) and Cruciferae (3%). The total pollen count from this trap was low (Appendix 4).

In the 1996 sampling period Eucalyptus was consistently represented with levels of 20% at the overhang (trap 6/1996) and 21% downslope (trap 5/1996). Levels of Asteraceae •140-

A \ / (topMoun)t MiddeChambern s 2.-K\

Mount Chambers 2. . Ptilotus obovatus (overhang) Midden . Poaceae spp , oieana decurrens . Acacia vicforiae TV . Ptilotus obovatus w ^ /. Atriplex spp

FOLIAGE COVER y p0LLENTRflP < >'t. ^r ROCKY OUTCROP * g'.'o^ El SOIL AND GRAVEL • 11-15% p] GRAVEL AND BOULDER • 16-20% 1^ TALUS SLOPE 0 l0 >K • > 20% m«1r«s

Trees Shrubs Herbs

J Jf Sf _4> &# J& .. sfjt& ;f.

in 1995 out 1995 in 1996 out 1996 Grasses onrm J e&J?•• r

I I I II Ll—I in 1995 out 1995 in 1996 0 10% out 1996

Summary Diagram & & «? ^ •

in 1995 out 1995 in 1996 out 1996

Figure 5.22: Pollen diagram of 1995 and 1996 traps (3 and 4) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Mount Chambers 2, Mount Chambers Gorge. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.16. -141-

ranged from 9% in trap 6/1996 to 4% in trap 5/1996, and Chenopodiaceae/ Amaranthaceae (24%) was more abundant in trap 6/1996 (Figure 5.23). Fabaceae (5%) and Sapindaceae (3%) were consistently represented in both traps. There was 4% Other Myrtaceae A in trap 6/1996 and 14% Other Myrtaceae C in trap 5/1996. The levels of Poaceae and Cyperaceae (5%) were more abundant in trap 6/1996 and herbaceous taxa were more diverse downslope in trap 5/1996 (Figure 5.23).

The signal from chenopods/amaranths was less abundant inside Chambers Gorge. In 1995, grasses were the most abundant signal downslope of CGI followed by trees and chenopods/amaranths. In 1996 trees increased and herbs and chenopods/amaranths decreased downslope. Upslope there was a stronger signal of herbs and chenopod/amaranths and less abundant trees (Figure 5.23).

Chambers Gorge 1 Vegetation CGI midden site was located at the top of a loose talus slope that extended down to Chambers Creek at the base. On stable sections of slope there were Eremophila freelingii shrubs and a ground cover dominated by Ptilotus obovatus, Sida petrophila, Solanum petrophilum, Stipa spp and Enneapogon spp.. Along Chambers Creek were Melaleuca glomerata and Eucalyptus camaldulensis trees and Abutilon leucopetalum on the floodplain (Plate 5.12).

Sixteen taxa recorded in the pollen were not in the vegetation cover upslope near the overhang and 15 taxa downslope in Chambers Creek. These included herbaceous and shrub taxa that are present in other sections of the gorge (Table 5.17).

Chambers Gorge MD3 Traps Levels of Eucalyptus were abundant inside (trap 7/1995) and outside the cave (trap 8/1995) ranging from 51% to 44% respectively. Melaleuca was consistently represented at levels between 12%-18% and less abundant Cupressaceae (2%-4%) and Casuarinaceae (1%) (Figure 5.24). Chenopodiaceae/Amaranthaceae ranged from 12% in trap 7/1996 to 15% in trap 8/1995 and these were the lowest recorded levels for this taxon at Mount Chambers Gorge. Asteraceae was consistently represented with levels between 5%-6% and there were other less abundant herbaceous taxa present in both traps (Figure 5.24). -142-

*> K < Chambers Gorge 1. Midden \^ •* stipa spp J / £nneapogon spp.

Ij^ %. £remophi/o freeJingii * >>-v Ptilotus obovatus »?^l / Sida petrophila '2y» Solanum petrophilum o"c\

-)fl.0«O °'Ps,

% SPECIES PRESENT FOLIAGE COVER (•«• leu) < IV. * trjelateuca giomenata T POLLEN TRAP • 1 - 5% Abuti/on /eucopetulum "< ROCKY OUTCROP • 6 - 10% 1 £ucal 20% a TALUS SLOPE Sv.i'.v^--. m.fr.t

Trees Shrubs Herbs

/s • ,>0 ^ ///,* ^.* * .# r ^ s d»^- cf * n i out 1995 in 1996 out 1996

rrrcrrrrrrrPFHerbs Grasses F

^^ «?ft? C* V <# _L| l| l| I I—L |—I—Lj—I—I out 1995 in 1996 out 1996 rFff 0 10% Summary Diagram

out 1995 in 1996 out 1996

Figure 5.23: Pollen diagram of 1995 and 1996 traps (5 and 6) from inside and outside the midden cave site and vegetation present at Chambers Gorge 1, Mount Chambers Gorge. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.16. -143-

Plate 5.11: Vegetation (downslope from MCI) outside MC2 cave site on the south western face of Mount Chambers.

Plate 5.12: Sparse vegetation including Ptilotus obovatus and Eremophila freelingii on the scree slope outside CGI midden site. -144-

Pollen Trap 7/1996 (inside the cave) recorded 55% Eucalyptus, 12% Chenopodiaceae/ Amaranthaceae and 11% Melaleuca. There was 4% Convolvulaceae, 3% Asteraceae and lower levels of Liliaceae, Malvaceae and Thymelaceae (Figure 5.24).

Trees were the most abundant category inside and outside the cave in 1995 and 1996 followed by herbs and chenopods (Figure 5.24).

Chambers Gorge MD3 Vegetation Transects Ptilotus obovatus, Eremophila freelingii, Cymbopogon ambiguus and Stipa spp. were growing near the cave opening at the top of the slope. Dodonaea viscosa and Casuarina cristata were present near the base of the slope and Eucalyptus camaldulensis trees with Lysiana exocarpi in the foliage, were growing along the creek bed (Plate 5.13 and Figure 5.24).

There were 15 taxa recorded in the pollen but not the vegetation inside the cave and 14 taxa outside. Loranthaceae was only recorded in the vegetation and this taxon is documented as entomophilous that can under-represent the source plant (Boyd 1992) (Table 5.17).

5.5.3 Brachina Gorge Cave Traps and Vegetation Surveys

Brachina Gorge 1 Traps Chenopodiaceae/Amaranthaceae was consistently represented inside (trap 10/1995) and outside (trap 1/1995) the cave ranging from 16% - 19%. Asteraceae (24%), Eucalyptus (18%) and Poaceae/Cyperaceae (21%) were more abundant in trap 10/1995 (Figure 5.25). Levels of shrub taxa were higher outside the cave and included 13% Sapindaceae, 5% Fabaceae and relatively less abundant Acacia, Myoporaceae and Solanaceae. Herbaceous taxa including Liliaceae and Convolvulaceae were present at low levels in both traps (Figure 5.25).

There were high levels of Fabaceae (34%) and Convolvulaceae (23%) recorded outside the cave in 1996 (trap 1/1996). Chenopodiaceae/Amaranthaceae contributed 9%, there was 7% Myoporaceae and 5% Eucalyptus (Figure 5.25). Shrub taxa included Caesalpinaceae, Sapindaceae and Santalaceae and a variety of herbaceous taxa such as Liliaceae, Asteraceae and Cruciferae were all <5% of the signal (Figure 5.25). •145-

* Ptilotut obovatus £remophila freelingii Cumbopopon om&guus Chambers Gorgi 2 otipa spp (M03) Mlddtn T

% SPECIES RECOROED AS PRESENT AT THE SITE * Dodonaea viscosa (•»• mil Casuarina cristata T POLLEN TRAP £ucaluptus camaldu/ens/o with "C ROCKY OUTCROP Lusiana erocarpi in the foliage [S] BOULDER AND SOIL IS] TALUS SLOPE

Trees Shrubs Herbs J

G<$ rf <£> 4? ^VVVWw <' tf «? ^wvv «? in 1995 out 1995 0 10% in 1996

Figure 5.24: Pollen diagram of 1995 and 1996 traps (7 and 8) from inside and outside the midden cave site and vegetation species at Chambers Gorge 2 (MD3), Mount Chambers Gorge. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.16. -146-

Pollen Traps Trap 1/95 | Trap 4/95 Trap 3/95 Trap 7/95 Trap 8/95 Cupressaceae * Casuarinaceae * Eucalyptus * Melaleuca * Proteaceae * * Acacia * * Myoporaceae * * * Caesalpinaceae * * * Sapindaceae * Fabaceae * Lamiaceae * * Euphorbiaceae * Goodeniaceae * * * Solanaceae * * * Apiaceae * Liliaceae * * Malvaceae * * * * Thymelaceae * * Convolvulaceae * * Cruciferae * * Onagraceae * * Onagraceae (2 pore) * * Haloragaceae * * Boraginaceae * * Cyperaceae * * * Poaceae * Echium spp. * Unknown * * *

Table 5.16: Taxa recorded at levels of less than 1% in cave traps from midden sites on Mount Chambers and in Mount Chambers Gorge for 1995 and 1996 sampling periods (1995 this page and 1996 continued on the next page). -147-

Pollen Traps Trap 1/96 Trap 2/96 Trap 4/96 ! Trap 3/96 Trap 5/96 Trap 6/96 Trap 7/96 Cupressaceae * * * * Casuarinaceae * * * * Eucalyptus * Myrtaceae A * * Acacia * * * * * Sapindaceae * Myoporaceae * * * Caesalpinaceae * * * * Santalaceae * Euphorbiaceae * * Goodeniaceae * Solanaceae * * * * * Malvaceae * * Cruciferae * * * * Liliaceae * * * * Thymelaceae * * * Convolvulaceae * * Onagraceae * * Onagraceae (2 pore) * Boraginaceae * * * Haloragaceae * * Cyperaceae * * Poaceae * * Plantaginaceae * * Echium spp. * * Pinus spp. * Unknown * * *

Table 5.16 (continued): Taxa recorded at levels of less than 1% in cave traps from midden sites on Mount Chambers and in Mount Chambers Gorge for 1995 and 1996 sampling periods. -148-

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In 1995, herbs were most abundant both inside and outside the cave and levels of trees and grasses were higher inside. Signals of chenopods/amaranths were similar in each trap. In 1996, shrubs and herbs were dominant and other categories were less abundant compared to the previous sampling period (Figure 5.25). The level of chenopods, trees and grasses had dropped in 1996. During summer 1995 to early winter 1996 it was drier compared to the previous sampling period and this may have been reflected in the reduced pollen signals from these vegetation categories.

Vegetation transects at Brachina Gorge 1 A patch of Alectryon oleifolium tall shrubs contributed to 11% foliage cover outside the cave entrance, with a sparse ground cover of Chrysocephalum semicaluum and Olearia decurrens and more abundant Ptilotus obovatus and Sida petrophila (Figure 5.25). Dodonaea viscosa spp. angustissima shrubs were also recorded along the transect. On the lower sections of slope there was a decrease in the foliage cover of trees and shrubs with the addition of Acacia tetragonophylla and Senna artemisioides. An increase in ground cover species included Chrysocephalum semicaluum, Sida petrophila, Rhagodia parabolica and Olearia pimeleoides (Figure 5.25). Other species along the transect included Bursaria spinosa, Ptilotus obovatus, Pimelea microcephala, Abutilon leucopetalum, Solanum petrophilum, Enchylaena tomentosa and Zygophyllum aurantiacum. Cymbopogon ambiguus, Solanum ellipticum, Euphorbia drummondii and Enchylaena tomentosa were present along the transect at the base of the slope (Plate 5.14).

There were an additional 12 taxa recorded in the pollen traps inside and outside the cave that were not in the vegetation transects. Pittosporaceae, Thymelaceae and Zygophyllaceae were not recorded in the traps and the latter two are known to be under representative of source plants (Table 5.19).

Brachina Gorge 2/7 Traps Levels of Cupressaceae (40%) and Eucalyptus (8%) were more abundant inside the cave (trap 5/1995) compared to outside (trap 6/1995) (Figure 5.26). The representation of Apiaceae ranged from 36% in trap 6/1995 to 9% in trap 5/1995 whereas Asteraceae was more consistently represented with levels ranging from 10% inside and 7% outside the cave. Other herbaceous taxa less abundant outside the cave included Thymelaceae, Malvaceae and Onagraceae. Chenopodiaceae/Amaranthaceae was consistently represented in both traps with levels ranging from 11%-16% (Figure 5.26).

Eucalyptus (15%-20%) and Cupressaceae (2%-4%) were consistently represented inside (trap 5/1996) and outside (trap 6/1996) the cave in 1996. Levels of -150-

Chrusocephalum semicaluum Olearia decurrens Ptilotus obovatus Dodonaea viscosa ssp angustissima Sida petrophila Alectripn oleifo/iium Brachina I. Dodonaea viscosa spp. angustissima Olearia pimc/eoides Midden Cnrusocepha/um semicaluum Alectryon o/ejofoliuni Sida petrophila Olearia pimeleoides Pnapodia parabolica Pimelea microcepho/a Ptilotus obovatus FOLIAGE COV E» T POLLEN TRAP Dodonaea viscosa spp angustissima

• \t. <; -< ROCKY OUTCROP Cassinia laevis . i -5% CArusoccphalum semicaluum 6 - 10% • f>] SOIL AND GRAVEL Sursana spmosa • II - IS% • 16 -20% Sida petrophila • > 20% 0 10

Trees Shrubs Herbs //////4///4*' -& ^ A #/ / J|_l l_J II i| M ' ' i| i in 1995 out 1995 out 1996 FF Herbs Grasses

1 J .jf •J „<#

' ' 'I 1 in 1995 out 1995 out 1996 0 10% Summary Diagram

& m -*< 4>& *& V^ V & £>

in 1995 out 1995 out 1996

Figure 5.25: Pollen diagram of 1995 and 1996 traps (1 and 10) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Brachina 1, Brachina Gorge. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.18. -151-

Plate 5.13: Vegetation downslope of MD3 cave site at the eastern end of Mount Chambers Gorge including Eucalyptus camaldulensis, Dodonaea viscosa and ground cover including Ptilotus obovatus and Stipa spp.

Plate 5.14: Eremophila freelingii, Dodonaea viscosa and Alectryon oleifolium shrubs growing on the slope outside BR1 cave midden site. -152-

Chenopodiaceae/Amaranthaceae ranged from 25% inside to 9% outside and Cruciferae was 5% inside versus 34% outside the cave. Poaceae and Cyperaceae were similar in both traps (Figure 5.26).

In 1995, the levels of trees and herbs were most abundant inside the cave. Shrubs and chenopods/amaranths were similar in both traps. In 1996, the tree signal inside the cave was reduced and chenopods/amaranths had increased. The abundant herb signal was still recorded outside in 1996, trees had increased and chenopods/amaranths had decreased (Figure 5.26).

Vegetation Transects at Brachina Gorge 2 Dodonaea viscosa spp. angustissima, Hakea ednieana and Prostanthera striatiflora were present above the cave, with a sparse ground cover of Stipa spp., Sida petrophila , Ptilotus obovatus and Bursaria spinosa (Figure 5.26). Solanum ellipticum was also present along the transect. The foliage cover of ground species increased outside the cave entrance and the shrubs decreased. Additional species at this site included Eremophila latrobei, Pimelea microcephala, Zygophyllum aurantiacum and Enchylaena tomentosa. A ground cover of Zygophyllum aurantiacum and Senna artemisioides spp. artemisioides (affinities to spp. petoilaris) shrubs, were dominant at the base of the slope. Other shrubs at this elevation included Dodonaea viscosa spp. angustissima and Acacia ligulata with a ground cover of Pimelea microcephala, Sida petrophila and Stipa spp.(Plate 5.15).

There were an additional 12 taxa recorded in the pollen and not the vegetation inside the cave and 14 taxa from the trap outside. Pittosporaceae and Proteaceae were recorded in the vegetation but not the pollen in both sampling periods and Lamiaceae, Thymelaceae and Malvaceae were only recorded in 1995 (Table 5.19).

Brachina Gorge 3/4 Traps Eucalyptus was not consistently represented at this site with levels ranging from 24% in trap 8/1996 (outside the cave) to <2% in trap 7/1996 (inside). Cupressaceae (6%) and Casuarinaceae (5%) levels were also more abundant outside the cave (Figure 5.27). There was 11% Pittosporaceae in trap 8. Asteraceae was more consistently represented (2% - 3%) inside and outside respectively. Chenopodiaceae/Amaranthaceae (12%) was more abundant in trap 8/1996 whereas Poaceae and Cyperaceae were similar in both traps (Figure 5.27). Exotic taxon Echium spp. contributed to 74% of the signal inside the cave compared to 2% outside.

Exotic pollen was abundant inside the cave and other vegetation categories were less than 10% of the pollen sum. Levels of tree pollen were higher outside the cave and the -153-

. Hakea ednieana . Prostonfnera strtafif/cva , Burjoria spvhoaa . Sida petrophila ~T~^r~Tri . Ptilotus obovatus . Stipa spp. Brachina 7 —- . Dodonaea viscosa spp. anpusfissimo Midden x)L \Tk T(*I^^r ( . Dodonaea viscosa spp anyusiissima ^^TjJ , Prostanthera striatiflora s^^^T , Bursona spinasa , Pimelea microcephala Brachina 2. . Sida petrophila Midden TS* . Ptilotus obovatus 'L'S^s. . Hakea ednieana ••'••:'L\ . Acacia victoriae '•'•.:;?Sw . Senna arttmisiodies spp petio/aris •L^>~v^ • Bursana spmosa ''^viLl^VvSv • Zitgophullum aurantiacum '"••:'^.\ . Solanum ellipticum \V:';\. , marrubium vulpare FOLIAGE COVER • < IV. T POLLEN TRAP • 1-3% \ R0CK1 OUTCROP • 6 - 10% so • 11-15% E3 "- 4ND GRAVEL • 16-20% •:'-';\ 0 10 • > 20% "'* 1 ,., ' m.tr.i

Trees Shrubs Herbs SS s»y/f/s9> # -&.

cr cf 1 ^ in 1995 out 1995 in 1996 out 1996 Herbs Grassews // // /y/y/*

in 1995 out 1995 in 1996 out 1996 0 10%

Summary Diagram

«5* tf 4*^ $f «>* ? J*- A&v is° ^ ^ & # C^

in 1995 out 1995 in 1996 out 1996

Figure 5.26: Pollen diagram of 1995 and 1996 traps (5 and 6) from inside and outside the midden cave site and foliage cover of species along transects parallel to the slope contour at Brachina 2/7, Brachina Gorge. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.18. -154-

abundance of herbs, grasses and chenopods/amaranths were similar to each other. There were lower levels of shrubs both inside and outside the cave (Figure 5.27).

Vegetation at Brachina Gorge 3/4 At BR3/BR4 midden site, the slope was too steep for transects. Eucalyptus camaldulensis, Exocarp aphyllus, Acacia continua and Acacia tetragonophylla were present at the base of the slope. On the rocky slope rising from Brachina Creek to the cave there were Dodonaea viscosa shrubs growing out of pockets of soil and a sparse ground cover of Sida petrophila, Ptilotus obovatus and Stipa spp. Casuarina cristata and a small Hakea ednieana tree were present on a rocky talus slope, opposite the cave entrance (Plate 5.16).

There were an additional 10 taxa recorded in the pollen but not the vegetation inside the cave and 11 taxa outside (Table 5.19). Proteaceae and Malvaceae were only recorded in the vegetation surveys and are known to be poorly dispersed pollen (Table 5.12).

Comparison of taxa in pollen spectra and the vegetation cover at cave sites SI values calculated for cave pollen traps at each midden site illustrated differences in the modern pollen spectra from this local investigation. Generally, there was a higher degree of similarity between taxa in the modern pollen rain and vegetation at cave sites compared to traps along west-east transects across the ranges. This is a reflection of capture of local taxa in vegetation on the slopes outside the cave in addition to the regional component. Local environmental features enhanced the capture of local and often under-represented taxa. The majority of sites from the Arkaroola-Mount Painter Sanctuary recorded SI values 0.4 and above (Table 5.20). Sites from steep rocky locations including North Well Creek 1 and Radium Creek 2 with a sparse vegetation cover on the slopes recorded lower SI values similar to those from the west-east transect traps. The SI value from site MCI on Mount Chambers was lower compared to the other midden sites (Table 5.21). This is largely a result of the sparse vegetation cover on the rocky slope outside the cave. At Brachina Gorge (Table 5.22), the degree of similarity between modern pollen and vegetation was similar to SI values recorded at some of the Arkaroola midden sites. -155-

# Sida petrophila Ptilotus obovatus Stipa spp. Casuarina cristata ( I I reel Hakea ednieana (I shrub) Dodonaea viscosa Brachina 4 Midden

& Eucaluptus camaldulensis 1 Jff SPECIES PRESENT ON SLOPE £xocarpus aphul/us (i.t ttitl Acacia confinua Y POLLEN TRAP Acacia tetragonophul/a < ROCKY OUTCROP £3 SOIL AND GRAVEL

Trees Shrubs Herbs *.•

'''' in 1996 out 1996 0 10%

Qrasses Summary Diagram

**

Figure 5.27: Pollen diagram of 1995 and 1996 traps (7 and 8) from inside and outside the midden cave site and vegetation species at Brachina 3/4, Brachina Gorge. Pollen sum equals total pollen counted. Taxa recorded at levels of less than 1% are listed in Table 5.18. -156-

Follen Traps Trap 10/95 Trap 1/95 Trap 5/95 ! Trap 6/95 j Casuarinaceae * * Melaleuca * * Myrtaceae A * * Acacia * * Sapindaceae * Myoporaceae * * Proteaceae * Caesalpinaceae * Lamiaceae * * * Santalaceae * Euphorbiaceae * * * Goodeniaceae * * Solanaceae * * Thymelaceae * Malvaceae * * Convolvulaceae * * Liliaceae * * * Haloragaceae * Zygophyllaceae * Onagraceae * Echium spp. * * Plantaginaceae * Pinus spp. *

Pollen Traps Trap 1/96 Trap 5/96 Trap 6/96 Trap 7/96 Trap 8/96 Cuppressaceae * Casuarinaceae * * * Melaleuca * Proteaceae * Acacia * * * * Sapindaceae * * Lamiaceae * Caesalpinaceae * * Solanaceae * * Santalaceae * Euphorbiaceae * Epacridaceae * Apiaceae * Thymelaceae * Liliaceae * * * Convolvulaceae * Malvaceae * Haloragaceae * * Onagraceae * * * * Zygophyllaceae * Lythraceae * Boraginaceae * * Plantaginaceae * * Cyperaceae * Poaceae * Echium spp. * Unknown * *

Table 5.18: Taxa recorded at levels of less than 1% in cave traps from midden sites in Brachina Gorge for 1995 and 1996 sampling periods. -157- Trap Site Brachina 1 I Brachina 2/7 I Brachina 3/4 Year 1995 1995 1996 | 1995 1995; 1996 1996 1996 1996 Location in out out in out | in out in out Trap Number 10! 1 1 5 6 5 ei 7 8 i ; I Cupressaceae * * * i * ; * * I * * Casuarinaceae I * * * : i* i I* I. Eucalyptus j * * * ; * * * I* I j* :• . Melaleuca * I* * i* I* ! ! ,* I Other Myrtaceae * i * i * j i j ; i Acacia i* * '• !* !* . * !* :• : * :* .« Proteaceae * i» ! i |« i i« * * j * Sapindaceae * * !• !• i* * I *

* , * j * i Myoporaceae * * * * i (• | I Caesalpinaceae * !• « ; !• I* i Solanaceae * * * • * I * * I * ; _ : * I Ii. i Euphorbiaceae | * * I* I* I* ! I ! I Goodeniaceae ! * !• i* ! Asteraceae j* i* !• I* I* I* I |* !* | Apiaceae ! * I* : '* I* * I* I •' i* Fabaceae i* ;* * j* |* * ;* i* :* Lamiaceae I* i I* j* * ;• Epacridaceae • I1* j Liliaceae i* i* !* ;* ;**!*; i* Thymelaceae •;* • !* !* 1* • Malvaceae !* i I* ;•'*!* i !• ! ;• Convolvulaceae 1* j* i ' !* * ! i* |* Haloragaceae ill 1* !* i* Santalaceae . i * i i * 1 \ '•• ! * ! • Onagraceae . 1 * ;* * i* :* * Lythraceae ; * i Cruciferae !* i* ';* * ••* * i* ••* Zygophyllaceae 1 • !* i* •• Pittosporaceae 1 > : :• Chenopodiaceae :* ,*;*.* • * * j* Cyperaceae ;* i* ;* i* Poaceae !* \* !* •• •' 1* * i* • |* Exotics I* 1* 1* i* !* i* ill i i . , • , : .; i I | . i ' | ' : : I 1 I *= taxon in pollen traps i ' ! ' ' • \ •= taxon recorded in vegetation '

Table 5.19: Comparison of taxa recorded in the modern pollen and vegetation at midden cave sites in Brachina Gorge. Data was used to calculate a Sorenson Similarity Index for each site. -158-

H

.*-"

"^ • --'--' ,•- 9* W * ' ft' ^ t J **•* • 44 1

Plate 5.15: Shrubland dominated by Eremophila spp. and ground cover of Ptilotus obovatus and herbs at the western end of Brachina Gorge looking out from the entrance ofBR2/7 cave site.

Plate 5.16: Steep rocky slope and vegetation cover outside BR3/4 midden cave site. Casuarina cristata was growing outside the cave entrance. -159-

Cave Site 1995 1995 in out Haematite Hill 0.52 0.58 North Well Ckl 0.32 0.39 North Well Ck 2 0.48 0.56 Waterfall * 0.43 Arkaroola 1 0.58 0.57 Radium Ck 1 0.53 0.57 Radium Ck 2 0.25 0.3 Radium Ck 3 * * C^paminda Track 0.55 0.44 pollen trap not recovered

Table 5.20: SI values for a comparison between vegetation cover and modern pollen rain at cave traps from stick-nest rat midden sites in the Arkaroola-Mount Painter Sanctuary

Cave Site 1995 1995 in out Mount Chambers 1 0.28 * Mount Chambers 2 0.44 0.41 Chambers Gorge 1 * 0.53 Medlin Site 3 0.56 0.47 * pollen trap not recovered

Table 5.21: SI values for a comparison between vegetation cover and modern pollen rain at cave traps from stick-nest rat midden sites at Mount Chambers Gorge.

Cave Site 1995 1995 in out Brachina Gorge 1 0.6 0.46 Brachina Gorge 2 0.54 0.54 Brachina Gorge 3/4 * * * pollen trap not recovered

Table 5.22: SI values for a comparison between vegetation cover and modern pollen rain at cave traps from stick-nest rat midden sites at Brachina Gorge. -160-

5.6 Modern Pollen Spectra at Midden Cave Sites

Chi-Square Tests Paired traps (inside and outside the cave) for the 1995 and 1996 sampling periods from each study site were compared using a Chi-squared significance test. Pollen data was grouped into categories of trees, tall shrubs, shrubs, herbs grasses and chenopods/amaranths. From all pairs of traps collected from Arkaroola, Mount Chambers Gorge and Brachina Gorge, only two pairs were found to be not significantly different from each other at a 0.05 level of significance (Appendix 8). One pair was located at Oppaminda Creek in the Arkaroola-Mount Painter Sanctuary (Trap 21/1995 and Trap 22/1995) and the other, at the eastern end of Mount Chambers Gorge (Trap 7/1995 and Trap 8/1995). Overall, modem pollen deposited inside the caves varied from the composition outside. The local environmental context plays a major role in determining the pollen rain composition irrespective of the vegetation cover at the site. At the two sites where the composition of pollen rain was similar both inside and outside the cave, the inside trap was relatively close to the entrance of the cave. This may have a stronger influence in capturing similar pollen taxa being deposited outside the cave.

Pollen Representation Cave traps have recorded a large number of pollen taxa contributing to less than 1% of the pollen sum. Arkaroola traps have recorded Casuarinaceae, Acacia, Myoporaceae and Caesalpinaceae at these levels, and herbaceous taxa such as Zygophyllaceae, Haloragaceae and Onagraceae (Table 5.14) during the 1995 sampling period. In 1996, there were generally less taxa at levels of <1% but these included Acacia, Myoporaceae, Euphorbiaceae and Convolvulaceae (Table 5.14). At Mount Chambers Gorge, similar taxa were found, together with Malvaceae, Solanaceae and Cyperaceae in 1995, and Liliaceae, Casuarinaceae and Solanaceae during 1996 (Table 5.16). During 1995 at Brachina Gorge, levels of <1% for Liliaceae, Euphorbiaceae and Lamiaceae were recorded, while in 1996 there was <1% Acacia, Onagraceae and Casuarinaceae (Table 5.18). Attention has been given to poorly represented taxa, as they are commonly recorded in midden pollen at higher levels. This has important implications for using these taxa to reconstruct Holocene vegetation communities.

The representativeness of these taxa in the pollen spectra across the Flinders Ranges is summarised in Table 5.12. All of these taxa have been documented as under representing source plants and/or having poorly dispersed pollen. Casuarinaceae is an exception, thought to be a regionally dispersed taxon. Contributions of poorly dispersed taxa that are known to be present at the cave sites is an indication that the cave traps are detecting quite a strong local signal. -161-

Taxa recorded in the modern pollen, compared to taxa recorded in vegetation transects have been summarised in Table 5.15 at Arkaroola, Table 5.17 at Mount Chambers Gorge and Table 5.19 at Brachina Gorge. At Arkaroola cave sites, pollen taxa that have been recorded in the majority of traps are also present at the cave sites. Taxa including Sapindaceae, Myoporaceae, Caesalpinaceae, Malvaceae, Chenopodiaceae/Amaranthaceae and Poaceae are examples (Table 5.15). Signals from tree and herbaceous taxa suggest a regional and extra-local component and a local source from herbs that may not have been present during the 1995 sampling period when vegetation transects were completed. A similar trend was observed at Brachina Gorge. Taxa included Acacia, Sapindaceae, Solanaceae, Thymelaceae, Malvaceae, Chenopodiaceae/Amaranthaceae and Poaceae which corresponded to species that were recorded in transect studies outside cave sites (Table 5.19). Mount Chambers Gorge is different from the other two study sites in terms of the vegetation communities present at the cave sites. There was a sparse vegetation cover of primarily chenopod/amaranths on the south western face of Mount Chambers Gorge and the sheer talus slopes of the gorge walls restricted vegetation cover to riverine woodland patches along Chambers Creek and shrub and herbaceous plants on the slopes. There was a less diverse suite of taxa recorded in vegetation transects from Mount Chambers Gorge trap sites in contrast to sites at Brachina Gorge and Arkaroola (Table 5.17). However diversity in the modern pollen rain was comparable to signals at Arkaroola and Brachina Gorge, suggesting a larger input from regional windblown pollen and local herbaceous species that were not detected during 1995 when vegetation transects were completed.

In summary, differences in pollen assemblages between traps located inside and outside a cave site, while significant, were not predictable, irrespective of the local vegetation communities at the cave sites or whether the traps were located at Arkaroola, Mount Chambers Gorge or Brachina Gorge. The influence of topography and aspect at each cave site was strong so in effect there was a unique composition of modern pollen at each cave site. Generally, the pollen rain was composed of a stronger local signal in addition to inputs from extra-local/regional taxa at some of the more exposed sites. This was evident when comparing pollen signals to the taxa represented in the vegetation transects at various altitudes from the caves.

5.7 Comparison Between Transect and Cave Trap Studies

It was apparent at some of the sites that tree pollen levels were more abundant in cave traps as opposed to open trap sites in the regional study. The source of this pollen is most likely regional and capture (in the traps) appears to be enhanced if traps are located at higher elevations on slopes. The degree of exposure of a cave site may also be an important factor as traps located near open caves (eg North Well Creek 2, Radium Creek -162-

2 and Mount Chambers) recorded higher levels of taxa such as Cupressaceae in the absence of large numbers of Callitris columellaris in the vegetation.

Some pollen taxa were consistently recorded in only one of the six month sampling periods during the trap study. Included in this group were Malvaceae, Proteaceae, Lamiaceae and Solanaceae. At other sites, some of these taxa were recorded in the vegetation but not the modem pollen. This is most likely a consequence of pollen production and dispersal characteristics of these taxa, and different responses from annual, ephemeral or perennial species to variable rainfall during the sampling periods. These taxa were not always present at the time of vegetation sampling. It was observed that these taxa were more often recorded at cave sites as opposed to the open sites along the west-east transects, emphasising a stronger local pollen signal collected from traps inside and outside caves.

This study provided insight into some spatial variability between midden study sites. Mount Chambers was the only area where Chenopodiaceae/Amaranthaceae signals were most abundant, a strong representation of both local and regional abundance inside and outside the caves. Levels may be exaggerated due to the location of cave sites (especially on the exposed south western face of Mount Chambers) and close proximity to extensive saltbush plains that flank the Ranges and Mount Chambers Gorge. Such high levels of Chenopodiaceae/ Amaranthaceae pollen are not recorded from other cave sites at Arkaroola and Brachina Gorge, or from traps along the west-east transect study. There were generally higher levels of herbaceous pollen taxa recorded in the cave traps compared to open trap sites in the transect study.

As understanding the characteristics of modem pollen rain provides the foundation for interpreting fossil pollen records from middens, two important questions need to be addressed. 1. How will a Flinders Ranges stick-nest rat pollen record differ from a regional playa lake record? 2. What will the collecting/nesting/excreting activities of stick- nest rats do to the pollen record on top of other variability that has been identified so far in this chapter.

1. Middens and lake cores clearly have different pollen catchments. Pollen analysis from sediment cores records a dominance of regional pollen types that can mask low pollen producers (usually entomophilous taxa) and, as a result, pollen spectra of some vegetation communities can not be distinguished. Luly's (1990) modem pollen study at Lake Tyrell indicated that the majority of modem pollen deposited at Lake Tyrell in semi-arid southeast Australia was wind blown, in contrast to humid environments where pollen is predominantly washed from the catchment or carried in creeks. It can thus be difficult in core records to interpret vegetation changes and identify communities from locations -163-

where dominant species are not wind pollinated (Anderson and VanDevender 1995) and when there is a range of species covered by each pollen type in a mosaic patterning of vegetation characteristic of semi-arid environments (Thompson and Kautz 1983). Composition of pollen in sediments can be affected by a change in facies, velocity and turbulence of water transported sediment and there can be differential destruction of grains after deposition by microbial digestion and oxidation (Thompson and Kautz 1983; Markgraf et al. 1997). In arid regions, differentially re entrained pollen from depositional surfaces has the potential to bias pollen percentages in fossil samples especially when primary pollen production is low (O'Rourke 1990).

Midden pollen can add taxa of regional significance that are not identified as locally occurring species at midden sites, in addition to recording local taxa (Anderson and VanDevender 1995; Markgraf et al. 1997). Airborne pollen contributed to middens comes directly from the air as well as adherence to plant macrofossils (O'Rourke 1991). Windblown pollen recruited into middens is dependent upon how open the site is, the length of midden formation and nature of surrounding vegetation (Thompson 1985). The pollen content of Neotoma middens also reflects the season of plant macrofossil collection and species composition of the plants that have been incorporated (O'Rourke 1991). There are additional transport mechanism for pollen capture within middens that contributes to further variability in the sources for pollen.

2. As a result of nesting/collecting/excreting activities of stick-nest rats, some pollen taxa will be recorded at higher levels compared to signals recorded in the modem pollen rain. Recruitment of pollen into middens was discussed in detail in section 2.2.4 of Chapter 2. It is expected that levels of taxa including Chenopodiaceae/Amaranthaceae, Asteraceae (common building material) and Malvaceae (food source) may be exaggerated in midden pollen in contrast to levels in the modem pollen spectra. Reworking of pollen within midden deposits also has implications for the pollen spectra in middens. This can result from rats moving through the structure or mixing of pollen rich material while middens are being constructed. These important factors are revisited in a later discussion in section 8.2.1 in Chapter 8.

5.8 Conclusions

The regional modem pollen rain has been examined from traps along west-east transects through the Flinders Ranges and a local scale investigation of pollen deposition inside and outside midden cave sites. Modem pollen has to be interpreted on the basis of the variable nature of semi-arid vegetation communities, seasonality of rainfall, response of plant species to climatic conditions and the environmental features of individual midden sites in terms of topography, aspect and exposure of caves/overhangs. -164-

There was a larger number of taxa recorded in the modem pollen rain, compared to the composition of taxa recorded at each trap site as evidenced by SI values. This is a result of the variable nature of semi-arid zone vegetation communities in different seasons and changing abundances of perennial, ephemeral and annual species.

The location of traps in relation to different environmental settings has an effect on the composition of modem pollen. Traps buried on the open plains flanking the ranges are subjected to more widespread regional and extra local pollen taxa compared to traps under canopied vegetation where pollen taxa appears to be filtered by vegetation cover at the site.

The hierarchical cluster analysis suggests that spatial variability in the regional modem pollen rain is large and it is not possible to distinguish between transects at Arkaroola, Mount Chambers Gorge and Brachina Gorge. There was clustering of trap sites located within similar vegetation communities from all three transect locations. The cluster analysis did indicate some temporal variability between the 1995 and 1996 sampling periods. Traps from the same sampling period tended to cluster together irrespective of whether the traps were located at Arkaroola, Mount Chambers Gorge or Brachina Gorge.

Chi-squared significance tests on cave pollen traps indicated that there were significant differences between pairs of traps recovered from the 1995 and 1996 sampling periods. Pollen assemblages recorded inside as opposed to outside cave environments were unpredictable from site to site. However, the abundance of locally occurring taxa appearing in the modem pollen spectra was greater at cave trap sites that resulted in a higher degree of similarity between vegetation cover and cave traps (as evidenced by SI values), in contrast to the degree of similarity between vegetation cover and traps along west-east transects.

Midden pollen records provide insight into the response of vegetation to environmental changes. The limitations of these types of deposits include atypical locations in rocky upland environments. There is a potential for high levels of variability from different middens from the same study site, as a result of the influence of different environmental characteristics of cave/overhang locations. -165-

Chapter 6: Stick-nest Rat Midden Palaeoecological Records from the Northern Flinders Ranges: Arkaroola-Mount Painter Sanctuary

6.1 Introduction

This chapter provides a systematic description of the palaeoecological records from stick nest rat middens: radiocarbon dates, fossil pollen and macrofossil assemblages. Records from sites in the Arkaroola-Mount Painter Sanctuary, northern Flinders Ranges are presented in this chapter. The following chapter presents records from Mount Chambers Gorge and Brachina Gorge in the central ranges. There is an extensive spatial coverage of middens that provide a series of well dated sites that can be combined into a regional study of Holocene environments and palaeoclimates.

6.2 Arkaroola Middens

Seven middens (site 1-7) were collected along North Well Creek and tributaries, Arkaroola Creek and Radium Creek (Figure 6.1). Two other midden sites were located south of Arkaroola Village. Site 8 was near the Devils Slide and site 9 was 10km south along a tributary of Oppaminda Creek (Figure 6.1).

Deposits at Haematite Hill (site 1), North Well Creek (site 2 and 3), Waterfall (site 4) and Radium Creek (site 5, 6 and 7) are frorri caves in the Proterozoic Mount Neil Granite Porphyry, a massive red weathering granite and the Lower Proterozoic Freeling Heights Quartzite of schizt, quartz and mineral laminations. Towards the west at Haematite Hill, younger units of the Upper Callana Beds outcrop and include the Woodnamoka Phyllite and Blue Mine Conglomerate (Jones 1996). Deposits at Arkaroola 1 (site 8) and Oppaminda Creek (site 9) were found in Proterozoic Blue Mine Conglomerate in the Upper Callana Beds and the Tapley Hill Formation (Umberatana Group) of finely laminated blue-grey silty shales (Mount Painter Province Geological Map Sheet). Tables 6.1 to 6.9 describe the characteristics of each midden site. Site numbers on Figure 6.1 correspond with those referred to in Tables 6.1 to 6.9.

6.2.1 Waterfall 1 (WF1) midden

Seeds from the midden at Waterfall site were submitted for Accelerator Mass Spectrometry (AMS) radiocarbon dating and reported as post 1950 (OZC624) (Table 6.10).

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Shrubs were represented by Sapindaceae, Fabaceae and Myoporaceae. Herbaceous taxa were not diverse, comprising Apiaceae (4%) and Cruciferae (<1%) and there was Poaceae (4%) and Cyperaceae (3%) (Figure 6.2).

There was no Leporillus spp. bone or scat material in the deposit, suggesting that the midden may not have been built by Leporillus. The majority of macrofossils were bone (Table 6.11), most likely having originated from an owl deposit. Identified bones included a species of rodent tail bone, scapula fragments and skull fragments. Miscellaneous bone fragments could not be identified. There was a small clump of grey rodent hair and some small feathers in the midden. The botanical remains were Acacia spp. seeds and unidentified insect fragments were abundant in the assemblage.

Midden Code Material Dated Laboratory Code Conventional Age (yrs BP) HH1 TOP faecal pellet OZC040 590 ± 45 HH1 BASE faecal pellet OZC042 770 ± 70 NWCK1 faecal pellet OZC048 560 ± 60 NWCK2 faecal pellet OZC617U2 3 670170 RC1 faecal pellet OZC046 4 350160 RC2 faecal pellet OZC618 2 430 ± 80 RC3 TOP leaf OZC042 420+55 RC3 BASE leaf OZC043 450 + 60 ARK1 leaf OZC044 470 ± 65 ARK1 faecal pellet OZC045 620 ± 55 OT1 leaf OZC616 880 ± 75 OT1 faecal pellet OZC621U1 880 ±150 WF seeds OZC624 modern

Table 6.10: AMS radiocarbon dates for middens from the Arkaroola-Mount Painter Sanctuary.

6.2.2 North Well Creek 1 (NWCK1) Midden

This is a Late Holocene midden dated at 900±50 yrs BP (OZC048) from faecal pellets (Table 6.10). The midden was located along North Well Creek in a rocky outcrop on the eastern side of the channel. -177-

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Leporillus sp. 0.44 Dodonaea viscosa Daucus glochidiatus insect fragments Arkaroola 1 TOP Sida petroplula 1.35 Mil Ptilotus obovatus 0.11 Cassinia laevis 0.12 Malvaceae sp. 0.91 Leporillus sp. 2.7 Malvaceae sp. Cassinia laevis flowers Arkaroola 1 MIDDLE iJ5 Leporillus sp. 1.43 Rodent sp. vertebrae (x2) Cassinia laevis Asteraceae buds insect fragments 4.84 Aikaioola 1 BASE Rodent sp. tail bone Sida petroplula 0.25 Rodent sp. leg bone Malvaceae sp. 0.45 Rodent sp. vertebrae and atlas bone Dodonaea 0.15 Mouse size tail bone Myoporaceae leaves 4* Rodent sp. foot bone (<0.01) Rodent bone inside a claw Myoporaceae fragments 0.06 Unidentifiable fragments 0.16 Leporillus sp. (adult) iV.ii Callitris columellaris pod (Jolcopteran head Haematite Hill 1 Leporillus sp. (juvenile) (2.07) Coleopteran Ml. 01 abdomen Leporillus sp. 0.i6 Ptilotus obovatus Casuarina cristata dadodes Daucus glochidiatus Diptera pupae cases Radium Creek 1 Sida petroplula 0.08 Bee Cassinia laevis terminal is<0.0 I Brachy come sp. <0.0l

Leporillus sp. (juvenile) 0.14 grey/brown tuft Liioria cwingi (brown tree frog) left & nghl ilia Acacia so. Daucus glochidiatus Coleoptera Radium Crcsk 2 Leporillus sp. (adult) 2.87 feathets Nepltrurus levis (knob-tailed gecko) Contus sp. 17* fragments J0.08 grey hair Leggadinaforresti (mouse) Eclmiaria sp. 1* )0.03 carapace, elytra, Leggadina forresii right Ml in a fragment of Sclerolaeiui sp. 1* •appendages maxilla Gryllotalpa spp. Fused left and right nasal bones {Notomys (mole cricket) claw fuscus or Pseudomys gouldii) Post cranial bone fragments Part top of skink skull 1 Lower molar tooth mouse Led upper fragment incisor (medium mouse) Pseudomys gouldii (medium size mouse) fused sacral vertebrae Pseudomys boland 2 left incisors Left fragment of incisor {Pseudomys gouldii or Notomys fuscus Phascogale calura fragment ofright maxilla of red-tailed phascogale - Leporillus spp. Leporillus apicalis lower right molar Mi UJ Radium Creek 3 TOP U.M (incomplete formal) Ptilotus obovatus 0.01 Chenopodium spp. stem 1* Cassinia laevis 1.7 fragments L apicalis left upper molar M2 (incomplete Cassinia laevis leaves 0.2 (lowers formed) Acacia spp. 0.01 Rodeni sp. lower incisor (medium size mouse) Myoporaceae 0.91 Rodent sp. embryonic molars (x2)

Leporillus spp. 0.76 P. gouldii right upper incisor (medium size Cassinia laevis i i da petroplula bud Radium Creek 3 mouse) Sida petrophila 0.5 Sclerolaena burr 1* Diptera pupae cases Acacia spp. 0.01 Senna artemisioides stems 0.02 BASE Myoporaceae sp. 0.02 Dodonaea angustissima 0.17 COT" Leporillus spp. 47.VH Ptilotus obovatus Oppaminda Track 1 Acacia sp. <0.01 Insect fragments Myoporaceae sp. 0.61 Unidentifiable fragments1.6 2 '• Table 6.11: Macrofossil assemblages for middens from the Arkaroola-Mount Painter Sanctuary. * indicates the number of macrofossils present where weights could not be determined. -179-

There was Eucalyptus (38%) and Melaleuca (35%) recorded in the midden and Other Myrtaceae (4%). There was less abundant Asteraceae (5%), Myoporaceae (2%) and Chenopodiaceae/Amaranthaceae (2%). Other herbaceous taxa included Haloragaceae, Geraniaceae and Convolvulaceae, and Poaceae (1%) and Cyperaceae (2%) were present (Figure 6.2).

There was a diverse macrofossil assemblage from this midden that included faecal pellets, hair, bone/teeth, leaves, seeds, fruits and insect fragments (Table 6.11). Faecal pellets included a mix of Petrogale xanthopus (rock wallaby), Trichosurus vulpecula (brush tail possum) and Leporillus spp.(stick-nest rat) and there were strands of a species of rodent hair. This midden was most likely Leporillus and other types of faecal pellets originated from animals using the same cave. There was a foot bone of a mouse sized rodent and the left incisor of Notomys sp. (possibly N. fuscus). Leaf fragments included Ptilotus obovatus (Amaranthaceae), Cassinia laevis (Asteraceae), Senna oligophylla (Caesalpinaceae) and Acacia tetragonophylla (Mimosaceae) and there were Daucus glochidiatus (Apiaceae) fruits. A variety of seeds and pods were preserved in this midden from the Poaceae and Caesalpinaceae families and included Senna artemisioides pods, Digitaria senguinalist, Danthonia frigida, Sclerolaena spp. muricata and Enneapogon nigricans, Cornus sp., Genista sp., Cynoglossum sp., Dianella sp. and Bromus. Insect fragments consisted of Coleoptera elytra and weevils.

The shrub and herb taxa represented by plant macrofossils were also present in the midden pollen. There was no Eucalyptus or Melaleuca macrofossils but high levels of this pollen.

6.2.3 North Well Creek 2 (NWCK2) Midden

This midden was located on the western side of North Well Creek channel and returned an AMS date of 3 670+70 yrs BP (OZC617U2) on faecal pellets (Table 6.10).

There was less abundant levels of Eucalyptus (11%) and Melaleuca (20%) compared to the record from NWCK1 while levels of Chenopodiaceae/Amaranthaceae (22%) and Asteraceae (30%) were more abundant (Figure 6.2). Poaceae and Cyperaceae contributed 2% and 4% respectively. Shrubs and herbs were represented at low levels.

Macrofossils were less abundant in this midden and included Leporillus spp. faecal pellets, bones and insect fragments. This was a Leporillus midden located in cave that may also have been used as a roosting place for owls. The bones included miscellaneous unidentifiable fragments (Medlin pers.comm. 1996), lizard foot bones and vertebra of Nephrurus millii (barking gecko). Insect macrofossils included Gryllotalpa spp. (mole -180-

cricket) mouth parts and some unidentified fragments (Table 6.11). There was a small portion of unidentifiable leaf fragments (0.0 lg).

6.2.4 Radium Creek 1 (RC1) Midden

This is a Mid-Holocene midden located at the junction of Radium and Arkaroola Creek. An AMS date on faecal pellets returned an age of 4 350+60 yrs BP (OZC046) (Table 6.10).

There were high levels of tree and shrub taxa that included Eucalyptus (10%), Melaleuca (5%), Myoporaceae (14%), Fabaceae (8% ) and Caesalpinaceae (5%) (Figure 6.1). Other herbaceous taxa were diverse, comprising high levels of Asteraceae (25%) and low levels (<5%) of Apiaceae, Malvaceae, Liliaceae and Convolvulaceae and both Chenopodiaceae/ Amaranthaceae and Cyperaceae registered 6%.

Leporillus spp. faecal pellets were part of the macrofossil inventory along with Diptera pupae cases and a specimen of Hemiptera. This was exclusively a Leporillus midden. Botanical macrofossils included leaf fragments of Ptilotus obovatus (Amaranthaceae), Sida petrophila (Malvaceae), Cassinia laevis (Asteraceae) and Brachycome sp. (Asteraceae). Two fruits of Daucus glochidiatus (Apiaceae) and Casuarina cristata (Casuarinaceae) cladodes were present (Table 6.11). Plant macrofossil genera were present in the pollen assemblage.

6.2.5 Radium Creek 2 (RC2) Midden

This midden was located in a rocky outcrop on the eastern face of Radium Creek channel and dated at 2 430+80 yrs BP (OZC618) (Table 6.10).

There were high levels oi Eucalyptus (31%) and Asteraceae (21%) in the midden. Less abundant taxa included Chenopodiaceae/Amaranthaceae (13%) and Melaleuca (7%) (Figure 6.2). There were low levels of shrub and herbaceous taxa, Poaceae (4%) and Cyperaceae (2%).

There was an abundant assemblage of bone material in this midden; faecal pellets, botanical remains, strands of hair from a species of rodent, feathers and insect fragments. This was a Leporillus midden located in a shelter that may have been used as a roosting site by owls. The faecal pellets were from juvenile Leporillus spp. Seeds included Acacia sp. (Mimosaceae), Cornus sp., Echinaria sp. (Poaceae) and Sclerolaena sp. and there were fruits (<0.01g) of Daucus glochidiatus (Apiaceae) (Table 6.11). The insect remains included fragments of Coleoptera elytra and appendages, and a Gryllotalpa spp. (mole cricket) claw. The assemblage of bone included a significant find of the right and left ilia -181-

of Litoria ewingii (whistling tree frog), the first known recording of this species in the Northern Flinders Ranges (G.Medlin pers.comm. 1996). This is a medium sized brown tree frog usually found on ground or low vegetation on the banks of pools or streams, breeding opportunistically after rain. This species is the only member of the Litoria complex found in South Australia and Western Victoria (Barker et al. 1995). The Australian fossil frog record is confined to isolated bones in caves, dry lake beds and fresh water limestones (Barker et al. 1995). Leporillus spp. middens can be regarded as a new source for the preservation of bone material for the Australian fossil frog record. It is possible that the frog bones originated from an owl pellet incorporated into the midden matrix.

There were also bones of Nephrurus levis (knob-tailed gecko) and part of the skull of a species of skink. The remaining bones were mouse or rodent size and included post cranial fragments of Leggadina forresti (mouse), the lower molar and a fragment of the left upper incisor of a medium sized mouse and fused sacral vertebrae of Pseudomys gouldii (mouse) (Table 6.11).

6.2.6 Oppaminda Track 1 (OT1) Midden

This is another Late Holocene midden located south of the Arkaroola Village. There were AMS dates on both leaf fragments and faecal pellets that returned ages of 880±75 yrs BP (OZC616) and 880+150 yrs BP (OZC621U1) respectively (Table 6.10).

This midden recorded the highest levels of Chenopodiaceae/Amaranthaceae (66%) pollen for any site in the Arkaroola-Mount Painter Sanctuary. There was Asteraceae (6%), Apiaceae (5%) and lower levels of other herbaceous taxa including Convolvulaceae, Boraginaceae and Echium spp. (Figure 6.2). Shrubs were represented by Fabaceae (4%), Myoporaceae (3%) and Solanaceae (2%) and there was a weaker signal from tree taxa that included Eucalyptus (2% ), Casuarinaceae (<1%) and Melaleuca (<1%).

The macrofossil inventory included a mix of unidentifiable leaf fragments and Ptilotus obovatus (Amaranthaceae), Acacia spp. and Myoporaceae spp. leaves. Casuarina cristata (Casuarinaceae) cladodes were also recovered and a variety of different sized twigs from Myoporaceae shrubs. Leporillus spp. faecal pellets, unidentified insect fragments and Diptera pupae cases where also present (Table 6.11). Herbs were absent in the macrofossil inventory but recorded in the midden pollen at low levels. This midden appeared to be exclusively Leporillus. -182-

6.2.7 Radium Creek 3 (RC3 TOP) and (RC3 BASE) Midden

This midden was sampled along naturally occurring fracture planes at the top and base of the deposit. Leaves from the top and base sections were dated at 420+55 yrs BP (OZC042) and 450+60 yrs BP (OZC043) respectively (Table 6.10).

The pollen record from RC3 TOP recorded lower levels of tree pollen compared to older middens in the Radium Creek suite (Figure 6.3). Shrub taxa were more abundant and included Sapindaceae (15%), Solanaceae (9%), Caesalpinaceae (7%) and Myoporaceae (6%). There were high levels of Apiaceae (21%) and Asteraceae (18%), and other herbaceous taxa recorded levels of less than 2%. Poaceae (6%) was more abundant than Cyperaceae and Chenopodiaceae/Amaranthaceae (3%) was less abundant compared to levels in other Radium Creek middens.

In RC3 BASE, there was Sapindaceae (12%), Myoporaceae (9%) and Caesalpinaceae (5%). There was an increase in levels of Asteraceae (35%) and decrease in Apiaceae (12%) compared to the top section (Figure 6.3). Levels of Cupressaceae (3%) and Eucalyptus (4%) were higher than records in the top sample of midden while levels of Chenopodiaceae/Amaranthaceae (4%) remained similar. Poaceae (4%) and Cyperaceae (2%) signals were less abundant compared to the top section.

Overall, levels of trees, shrubs etc (summary diagram in Figure 6.3) were similar in the top and base sections of the midden with only a change in the proportions of different taxa within each category.

The macrofossil assemblage from the top and base sections of midden is listed in Table 6.11. Both sections contained Leporillus sp. faecal pellets. The leaf fragments included Sida petrophila (Malvaceae), Acacia sp. (Mimosaceae) and Cassinia laevis (Asteraceae) from both sections, and Ptilotus obovatus (Amaranthaceae) from the top only. There were Acacia spp. pods and Daucus glochidiatus (Apiaceae) fruits in the top and Sclerolaena spp. burrs at the base of the midden. The most abundant taxa in the midden pollen are represented by plant macrofossils. Coleoptera fragments were recovered from the top of the midden and unidentifiable fragments and Diptera pupae cases in the base (Table 6.11). Bones from the top section of midden included the lower right (Ml) and upper left (M2) molar of Leporillus apicalis, a species of rodent (medium mouse size) lower incisor and two embryonic molars. The base sample contained the right upper incisor of Pseudomys gouldii (medium size mouse). This midden was definitely Leporillus. -183-

6.2.8 Haematite Hill 1 (HH1 TOP) and (HH1 BASE) Midden

This midden was located at the junction of Arkaroola Creek and North Well Creek. Faecal pellets from the top and base section of midden were AMS dated and returned ages of 590+45 yrs BP (OZC040) for the top and 770±70 yrs BP (OZC042) for the base sections (Table 6.10).

There was 15% Chenopodiaceae/Amaranthaceae pollen in the base sample, 16% Asteraceae and 12% Apiaceae. Trees were represented by Eucalyptus (9%), Cupressaceae (5%) and Melaleuca (4%) (Figure 6.4). Levels of shrub taxa included Fabaceae (7%), Myoporaceae (5%), Goodeniaceae (4%) and 3% of both Caesalpinaceae and Sapindaceae. Poaceae (3% ) and Cyperaceae (4%) were also recorded in this sample.

Levels of Asteraceae (24%) and Chenopodiaceae/Amaranthaceae (20%) were marginally higher in the top of midden. There was also an increase in the levels of Apiaceae (10%) and Convolvulaceae (4%). Eucalyptus (6%) was present in similar abundances but there was a drop in the levels of other tree taxa (Figure 6.4). Shrubs were represented by Myoporaceae (4%), 3% of both Caesalpinaceae and Goodeniaceae and Euphorbiaceae (2%).

There were abundant adult and juvenile Leporillus spp. faecal pellets (98.46g) in this nest/midden sample, twigs from woody shrubs (indeterminate species of the Sapindaceae and Myoporaceae) and Callitris columellaris (Cupressaceae) seed pods. Insect remains included abdomen fragments and Coleoptera elytra (Table 6.11). This deposit was a good example of a Leporillus stick nest.

6.2.9 Arkaroola 1 (ARK1 TOP), (ARK1 MIDDLE) and (ARK1 BASE) Midden

Leaf macrofossils and faecal pellets returned AMS ages of 470±65 yrs BP (OZC044) and 620±55yrs BP (OZC045) respectively (Table 6.10). The faecal pellets were sampled from within a nesting chamber in the centre of the midden and the leaves were from the base section.

There were high levels of Asteraceae (42%) and 15% Chenopodiaceae/Amaranthaceae pollen in the base sample. Shrub taxa included Myoporaceae (6%), Caesalpinaceae (4%) and lower levels of Solanaceae and. Acacia (Figure 6.5). Cupressaceae (3%) was the most abundant tree taxon, there was 4% Apiaceae and grasses were represented by Poaceae (4%) and Cyperaceae (2%). -184-

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Herbaceous taxa were dominant in the top sample of midden with 49% Asteraceae and 12% Liliaceae. There was 8% of both Chenopodiaceae/Amaranthaceae and Fabaceae and the record of Eucalyptus was similar to the middle section. Levels of Poaceae (3%) and Cyperaceae (2%) were similar to the signal recorded in the base. Levels of Asteraceae recovered and the signal of Chenopodiaceae/Amaranthaceae decreased.

The macrofossils recovered from ARK 1 included Leporillus spp faecal pellets, insect fragments and miscellaneous unidentifiable bone fragments (Table 6.11). Botanical macrofossils included Cassinia laevis (Asteraceae), Dodonaea viscosa (Sapindaceae), Sida petrophila (Malvaceae) and Ptilotus obovatus (Amaranthaceae) leaves and a mix of leaf fragments from ^determinate species of the Malvaceae. Daucus glochidiatus (Apiaceae) fruits were also preserved in the top of the midden. Lower levels of Asteraceae pollen in the middle section of midden occurred with a drop in the amount of Cassinia laevis macrofossils (0.02g). Daucus glochidiatus (Apiaceae) was recovered from the top section in the absence of Apiaceae pollen but not in the middle and base sections where pollen was present. The bone assemblage from an unknown rodent sp. was extracted from ARK1 BASE and included three vertebrae, tail bone, leg bone, atlas bone and a foot bone. There was a tail bone from a mouse size rodent sp. also present. This deposit was a good example of a Leporillus stick nest. The cave may have also been used by other animals including rock wallabies and owls.

6.3 Conclusion

The suite of middens from the Arkaroola-Mount Painter Sanctuary provided palaeoecological records for Mid-Holocene and Late Holocene environments in the northern Flinders Ranges. The next chapter presents pollen and macrofossil records from middens spanning the Early, Middle and Late Holocene periods at Mount Chambers Gorge and Brachina Gorge in the central Flinders Ranges. -188-

Chapter 7: Stick-nest Rat Midden Palaeoecological Records from the Central Flinders Ranges: Mount Chambers Gorge and Brachina Gorge

7.1 Introduction

This chapter presents descriptions of AMS chronologies, pollen records and macrofossil assemblages for stick-nest rat middens collected from Mount Chambers Gorge and Brachina Gorge. The oldest middens in this current study were located in the central Flinders Ranges.

7.2 Mount Chambers Gorge Middens

Stick-nest rat sites were found in both the Mount Chambers Ranges and Archeocyathid limestones at the eastern end of the gorge (Medlin 1993). They are in caves and fissures in Proterozoic sandstones, siltstones, shales with limestones and dolomites of the Tapley Hill Formation (Umberatana Group) (Arrowie Geological Map Sheet). Mount Chambers (site 10) and (site 11), were collected from caves on the south western slopes of Mount Chambers (Figure 7.1). Chambers Gorge (site 12) and Medlin (site 13) were from caves at the eastern end of Chambers Gorge (Figure 7.1). Tables 7.1 to 7.4 describe the characteristics of each midden site. Site numbers on Figure 7.1 correspond with those referred to in Tables 7.1 to 7.4.

7.2.1 Mount Chambers 1 (MCI TOP) and (MCI BASE) Midden

Faecal pellets from the centre of this midden returned an AMS date of 1 280+70 yrs BP (OZC620) (Table 7.5).

High levels of Chenopodiaceae/Amaranthaceae (90%) were recorded in the base of this midden. There was 1% Caesalpinaceae, Fabaceae and Sapindaceae and <1% Myoporaceae. Apiaceae and Asteraceae were 3% and 2% respectively while Cupressaceae, Casuarinaceae and Eucalyptus levels were all less than 1% (Figure 7.2).

High levels of Chenopodiaceae/Amaranthaceae (86%) remained in the top sample of MCI. There was 4% Apiaceae, 3% Asteraceae and other herbaceous taxa were <1%. Shrub pollen included Sapindaceae (2%), Myoporaceae (1%), Solanaceae (1%) and Caesalpinaceae (1%) (Figure 7.2). Eucalyptus, Cupressaceae, Casuarinaceae and Melaleuca were present at levels of <1%. The large number of pollen taxa consistently recorded at levels of less than 1% are listed in Table 7.6. Figure 7.1: Location of stick-nest rat midden sites at Mount Chambers Gorge. Site numbers correspond to those referred to in the midden descriptions in Table 7.1 through to Table 7.4. -190-

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Macrofossils (Table 7.7) included Leporillus spp. faecal pellets, insect fragments and pupae cases. The botanical remains were not abundant and included unidentifiable leaf fragments (0.2lg) and Sida petrophila (Malvaceae) leaves (0.17g). This deposit was a good example of a Leporillus stick nest.

7.2.2 Mount Chambers 1 (MCI EXPT) and (MCI EXPB) Midden

This Leporillus stick nest was located at the front of the cave housing MCI midden. An AMS date on faecal pellets from the midden returned an age of 410±65 yrs BP (OZC625) (Table 7.5).

In the base sample of midden there was 86% Chenopodiaceae/Amaranthaceae pollen. Asteraceae (5%) was the most abundant herbaceous taxon, and lower levels of Apiaceae, Malvaceae, Convolvulaceae, Liliaceae and Cruciferae were present. Shrub pollen taxa included 2% Fabaceae, Caesalpinaceae (1%), Myoporaceae (1%) and Solanaceae (1%) (Figure 7.2).

Chenopodiaceae/Amaranthaceae levels remained high (89%) in the top. Levels of Asteraceae (4%) were unchanged and other herbaceous taxa were present at levels of less than 1%. The record of shrub pollen was similar to the base sample and included 1% Myoporaceae, Caesalpinaceae and Solanaceae (Figure 7.2).

There were Sida petrophila (Malvaceae) leaf fragments, an abundance of twigs from the Chenopodiaceae family and Leporillus spp. faecal pellets in the macrofossil assemblage (Table 7.7).

7.2.3 Mount Chambers 2 (MC2 TOP) and (MC2 OVERHANG) Midden

There were AMS dates on leaf material and faecal pellets from the top section of MC2. Faecal pellets returned an age of 230+105 yrs BP (OZC622U1) and leaf material was 385±140 yrs BP (OZC614). Leaf material from the middle section of the midden overhanging the ledge, (MC2 OVERHANG) returned an age of 590±50 yrs BP (OZC623) (Table 7.5).

There was 87% Chenopodiaceae/Amaranthaceae, 3% Asteraceae and 2% Myoporaceae in the top pollen sample of MC2 (Figure 7.3). Levels of tree taxa were <1% and included Eucalyptus and Casuarinaceae. There was also <1% of herbaceous taxa, comprising Malvaceae, Thymelaceae, Onagraceae and Convolvulaceae.

There was 35% of Chenopodiaceae/Amaranthaceae, 30% of Asteraceae and 9% of Fabaceae in MC2 OVERHANG sample. Low levels of Asteraceae appear to coincide -195-

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with high levels of Chenopodiaceae that has also been observed in some of the middens from the Arkaroola-Mount Painter Sanctuary. Shrub pollen taxa included Myoporaceae (4%), Caesalpinaceae (4%), Solanaceae (2%) and Sapindaceae (1%) (Figure 7.3). Eucalyptus pollen contributed 3% to the pollen signal and herbaceous taxa included Apiaceae (3%), Cruciferae (3%) and 2% Malvaceae.

This deposit was exclusively Leporillus. The macrofossils in MC2 TOP included Panicum sp., Setaria sp. and Eleusine sp. Seeds (Table 7.7). Sida petrophila (Malvaceae), Ptilotus obovatus (Amaranthaceae), and an indeterminate number of species from the Myoporaceae and Malvaceae leaf fragments were also preserved. A large bulk of the midden was composed of small twigs and sediment that was not indurated with amberat. Insect fragments, Coleoptera thorax fragments and Diptera (?) pupae cases were recovered and there was an abundance of Leporillus spp. faecal pellets within the midden matrix (Table 7.7).

Macrofossils in MC2 OVERHANG included Daucus glochidiatus fruits, Ptilotus obovatus and Sida petrophila leaves. There were Diptera pupae cases, insect fragments and Leporillus spp. faecal pellets (Table 7.7).

7.2.4 Chambers Gorge 1 (CGI) Midden

This midden was most likely not exclusively Leporillus spp. as there were Petrogale xanthopus (yellow footed rock wallaby) faecal pellets identified in the midden matrix in addition to Leporillus pellets. Leporillus spp. faecal pellets returned an age of 470+130 yrs BP (OZC619) (Table 7.5).

There was 82% Eucalyptus pollen recorded in this midden, 2% Melaleuca and 3% Cyperaceae. Levels of Chenopodiaceae/Amaranthaceae (4%) were much lower than signals from Late Holocene middens located on Mount Chambers. There was 2% Asteraceae while levels of other herbaceous taxa were lower (Figure 7.4). Shrub taxa included 1% of both Myoporaceae and Caesalpinaceae and <1% Sapindaceae and Solanaceae.

The macrofossil inventory (Table 7.7) was not abundant as the midden was composed of amberat. Faecal pellets of Petrogale xanthopus (yellow footed rock wallaby) were recovered and identified on the basis of size, colour and coarseness of chewed material inside the pellet (Medlin pers.comm 1996). Leporillus spp. faecal pellets were also part of the inventory. Botanical macrofossils included Enneapogon polyphyllus and E. cylindric seeds and some unidentifiable leaf fragments. -198-

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7.2.5 Chambers Gorge 2 (MD3) Midden

Faecal pellets AMS dated at 7 260+100 yrs BP (OZC615U2) (Table 7.5) make this the oldest midden analysed in this current research.

There was 48% Chenopodiaceae/Amaranthaceae in the sample and lower levels of Eucalyptus (8%), Fabaceae (7%), Asteraceae (7%) and Myoporaceae (6%) (Figure 7.4). There was 4% Sapindaceae, 3% Melaleuca and 2% Caesalpinaceae. Herbaceous taxa were diverse and low levels were recorded of Onagraceae, Convolvulaceae, Thymelaceae and Malvaceae.

There was a variety of bone material in the midden possibly originating from an owl pellet that was incorporated into the stick-nest rat midden (Table 7.7). This was mostly species of rodent including a fragment of a leg bone (most probably the distal and proximal ends of the tibia), part of the left blade of the pelvis from a rodent similar in size to Pseudomys australis (plains rat), and the proximal end of a femur from a species of rodent. A proximal fragment of a right femur of possibly Leporillus apicalis (on the basis of size) was recovered as well as a broken-off keel of a sacral vertebra and the distal part of the pelvis of a medium sized rodent species. The other bones in the midden included a foot bone from a small marsupial species. There were unidentifiable leaf fragments (0.05g), leaf fragments from an indeterminate number of species from the Malvaceae, small twigs, insect fragments (0.02g) and Leporillus spp. faecal pellets (2.23g).

Midden Code Material Dated Laboratory Code Conventional Age (yrs BP)

MCI Centre faecal pellet OZC620 1280 + 70 MCI exposed base faecal pellet OZC625 410 + 65 MC2 TOP leaf OZC614 385 ±140 MC2 TOP faecal pellet OZC622U1 230 ±105 MC2 overhang leaf OZC623 590 ± 50 CGI faecal pellet OZC619 470 ±130 | MD3 faecal pellet OZC615U2 7 260 ±100

Table 7.5: AMS radiocarbon dates for middens from Mount Chambers Gorge. -201-

7.3 Brachina Gorge Middens

Brachina 1 (site 14), Brachina 2 and 7 (site 15) and Brachina 3 and 4 (site 16) were located at the western end of the gorge in caves in Lower Cambrian Wilkawillina Limestone (Figure 7.5). Tables 7.8 to 7.11 describe the characteristics of each midden site. Site numbers on Figure 7.5 correspond with those referred to in Tables 7.8 to 7.11. Middens were larger than middens from Mount Chambers Gorge and Arkaroola and were sub-sampled along natural fracture planes in the amberat, in order to investigate depositional history of middens.

7.3.1 Brachina Gorge 1 (BR1 TOP), (BR1 MIDDLE) and (BR1 BASE) Midden

Pollen was analysed from each section of midden and the top and base sections were AMS dated. Faecal pellets and leaves from the top and base sections were used to cross check ages of different types of macrofossils from the same sample and the results are presented in Table 7.4. There were AMS dates of 7 400+.90 yrs BP (BETA8108) on faecal pellets and 6 800+.70 yrs BP (OZB292) on leaf macrofossils for BR1 TOP. Ages from the base were 7 310±90 yrs BP (BETA81090) on faecal pellets and 6 710±80 yrs BP (OZB291) on leaf macrofossils (OZB291) (Table 7.12). These AMS dates and the implication for investigating depositional histories are discussed in the following chapter.

There was 19% Chenopodiaceae/Amaranthaceae pollen in the base of the midden, 15% Myoporaceae, 8% Caesalpinaceae and 5% Pittosporaceae. There was 10% Asteraceae and levels of other herbaceous taxa including Liliaceae, Cruciferae and Convolvulaceae were <5%. Eucalyptus (4%), Cupressaceae, Casuarinaceae and Melaleuca represented low levels of tree taxa in the base of the midden (Figure 7.6). A low signal of Polygalaceae is recognised as contamination.

The levels of Chenopodiaceae/Amaranthaceae (33%) and Asteraceae (28%) increased in the middle section of midden. Levels of Eucalyptus (8%) also increased in this sample while Casuarinaceae, Melaleuca and Cupressaceae remained low. There was a drop in the levels of shrub taxa while herbaceous taxa included 6% Apiaceae, 4% Liliaceae, and lower levels of Malvaceae and Lythraceae. The levels of Poaceae (4%) and Cyperaceae (4%) increased in the middle section. A low level of Boraginaceae pollen is attributed to contamination in the field sampling of this section of the midden.

There were high levels of Eucalyptus (19%) at the top of the midden and the signal from Chenopodiaceae/Amaranthaceae (27%) was similar to other sections. Asteraceae decreased to levels similar in the base section (9%). Shrub taxa included 8% Figure 7.5: Location of stick-nest rat midden sites at Brachina Gorge. Site numbers correspond to those referred to in the midden descriptions in Table 7.8 through to Table 7.11. -203-

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Myoporaceae, 6% Caesalpinaceae and 4% Fabaceae. There were low levels of Poaceae (2%) and Cyperaceae (1%) at the top of the midden (Figure 7.6).

Adult and juvenile Leporillus spp. faecal pellets were present throughout the midden (Table 7.14). There was an abundance of bone material including remains of large left and right upper incisors of Leporillus apicalis. The reminder of bone was derived from small mouse lumbar vertebrae and incisor fragments. Rodent species remains included thoracic, lumbar, sacral vertebra and part of a scapula of Notomys spp. (hopping mouse). The botanical macrofossils included Sida petrophila leaf fragments (Malvaceae) and other Malvaceae species. There were seeds from families including Poaceae, Caesalpinaceae and Sapindaceae. Other seeds were identified to genus and included Ranunculus spp., Galium spp., Cornus spp., Dodonaea viscosa and Cassinia laevis. Insect fragments were not identified but there were Coleoptera elytra in the assemblage. Macrofossils were less abundant from the middle section of this midden (Table 7.14) and included Acacia spp. leaf fragments and unidentified leaf fragments. There were leaf fragments of Cassinia laevis (Asteraceae), Sida petrophila (Malvaceae), Acacia spp. (Mimosaceae) and Dodonaea viscosa (Sapindaceae) in the sample from the base of the midden. Bone material included vertebrae and tail bone from a rodent species and the Ml and M2 left dentary, and M2 of the right maxilla of Notomys sp.(hopping mouse).

Taxa represented by plant macrofossils were recorded in the midden pollen from all sections of BR1 and ranged from trees to shrubs and herbs. Casuarina cladodes and Acacia leaves were recognisable macrofossils but the corresponding levels in the pollen were generally low.

7.3.2 Brachina Gorge 7 (BR7 TOP), (BR7 MIDDLE), (BR7 BASE) and Brachina Gorge 2 (BR2) Midden

An AMS date on faecal pellets from the base of BR7 returned an age of 5 790±80 yrs BP (OZC047). BR2 was located in the same cave as BR7 with an AMS date on faecal pellets of 5 020±70 yrs BP (OZC049U1) (Table 7.12).

Chenopodiaceae/Amaranthaceae (14%), Eucalyptus (13%), Fabaceae (9%) and Convolvulaceae (7%) were the most abundant pollen taxa in the base of BR7. Lower levels of tree taxa included Cupressaceae, Casuarinaceae and Melaleuca. There was 7% Asteraceae and other herbaceous taxa included Cruciferae (8%), Liliaceae and Boraginaceae (attributed to contamination due to the low level) (Figure 7.7). -208-

"46

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The levels of Chenopodiaceae/Amaranthaceae (26%) and Asteraceae (29%) increased in the middle section of BR7. Eucalyptus (6%) was less abundant and there was a continuation of lower levels of Cupressaceae, Melaleuca and Casuarinaceae. There was a high diversity of shrub taxa represented by low levels of Myoporaceae, Caesalpinaceae, Sapindaceae and Solanaceae. Levels of Poaceae (2%) and Cyperaceae (6%) had increased.

At the top of the midden there was a drop in the representation of Chenopodiaceae/Amaranthaceae (15%) and an increase in Cyperaceae (11%). Eucalyptus (9%) was the most abundant tree taxon at levels similar to the middle section of midden and levels of shrub taxa had increased with 7% Myoporaceae, 7% Fabaceae, Solanaceae (3%), Caesalpinaceae (3%) and Euphorbiaceae (3%). Asteraceae (15%) decreased and there were consistently lower levels of other herbaceous taxa such as Malvaceae, Convolvulaceae, Cruciferae, Haloragaceae and Thymelaceae (Figure 7.7). Midden Code Material Dated Laboratory Code Conventional age (yrs BP) BR1 TOP faecal pellet BET A8108 7 400 ± 90 BR1 TOP leaf OZB292 6 800 ± 70 BR1 BASE faecal pellet BETA81090 7 310 ±90 BR1BASE leaf OZB291 6 710180 BR2 MIDDLE faecal pellet OZC049U1 5 020 ± 70 BR7 BASE faecal pellet OZC047 5 790 ± 80 BR3 TOP faecal pellet OZB300 3 790 ±90 BR3 TOP leaf OZB299 3 570 ±120 BR3 BASE faecal pellet OZB293 3 640 ± 70 BR3 BASE leaf OZB298 3 690 ±180 BR4 (10cm) faecal pellet BET A81091 6 930 ±80 BR4 (10cm) leaf OZB295 6 290 ±190 BR4 (10cm) faecal pellet OZB294 5 690 ± 70 BR4 (20cm) leaf BETA87299 6 520 ±50 BR4 (30cm) leaf BETA87300 6 580 ±60 BR4 (40cm) leaf BETA87301 6 880 ±50 BR4 (50cm) leaf BETA87302 6 920 ± 50 BR4 (60cm) faecal pellet BETA81092 7 720 ±90 BR4 (60cm) faecal pellet OZB296 6 610 ±90 BR4 (60cm) leaf OZB297 6 650 ±80

Table 7.12: AMS radiocarbon dates for middens from Brachina Gorge. -211-

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The proportion of vegetation categories in different sections of the midden were not dissimilar but there was a change in the composition of taxa represented within the broad categories of trees, shrubs, herbs etc (see summary diagram Figure 7.7).

This deposit was a well indurated Leporillus midden. The macrofossil assemblage from BR7 TOP included Leporillus sp. faecal pellets and a variety of plant macrofossils that included leaf fragments of Sida petrophila (Malvaceae), Cassinia laevis (Asteraceae), Ptilotus obovatus (Amaranthaceae) and Dodonaea spp. (Sapindaceae) (Table 7.14). Leaf fragments from Dodonaea viscosa ssp. angustrissima (Sapindaceae) were also recovered from BR7 MIDDLE with the addition of Alectryon oleifolium spp. canescens (Sapindaceae), species of Asteraceae, Malvaceae and Sida petrophila (Malvaceae) leaf fragments. The insect assemblage included a mix of Coleoptera thorax and abdomen fragments. Diptera (?) pupae cases were also recovered from this section of midden. Macrofossils from the base section of midden were not as diverse or abundant. There was a variety of Cassinia laevis (Asteraceae), Dodonaea spp. (Sapindaceae), species of Malvaceae and Sida petrophila (Malvaceae) leaf fragments (Table 7.14).

BR2 midden recorded high levels of Chenopodiaceae/Amaranthaceae (68%). Shrub taxa included 4% Acacia, 3% Sapindaceae and 1% of both Myoporaceae and Caesalpinaceae. The most abundant herb was Cruciferae (6%) and other herbaceous taxa contributed to 14% of the pollen sum (Figure 7.7). There was no Cyperaceae in the midden, but there was 3% Poaceae. There was a weaker signal of trees and shrubs in BR2 compared to BR7, but a stronger signal of Chenopodiaceae. This may have resulted from BR2 being located closer to the cave opening and being more susceptible to capture of windblown taxa.

There were juvenile and adult Leporillus spp. faecal pellets in the midden matrix (Table 7.14). Bone material comprised fragments of skeletal material and teeth from a rodent species. A piece of rodent scapula was identified along with 12 vertebrae, foot and an ankle bone. There was also a premaxilla, tibia/femur and a piece of skull from a rodent species. A single left molar (Ml) of Leporillus apicalis was recovered and some unidentifiable bone fragments. Botanical remains included Cassinia laevis (Asteraceae) leaf fragments and Casuarina cristata (Casuarinaceae) cladodes. Unidentified insect fragments and Coleoptera elytra were sorted from the midden sample .

7.3.3 Brachina Gorge 3 (BR3 TOP), (BR3 MIDDLE) and (BR3 BASE) Midden

Samples from the top, middle and base sections of this Leporillus midden were collected, and macrofossils were extracted from the top and base sections for AMS dating. Leaves and faecal pellets from the top of the midden were 3 570±120 yrs BP -213-

(OZB299) and 3 790±90 yrs BP (OZB300) respectively. Leaves from the base of the midden were 3 690±180 yrs BP (OZB298) and faecal pellets returned an age of 3 640±70 yrs BP (OZB293) (Table 7.12). This was the youngest deposit in the suite of middens from Brachina Gorge.

There was 31% Chenopodiaceae/Amaranthaceae pollen at the base of BR3. Levels of herbaceous taxa included 11% Onagraceae, 7% Asteraceae, 4% Fabaceae and less abundant signals from Cruciferae and Liliaceae (Figure 7.8). There was 7% Eucalyptus and 4% of both Cyperaceae and Poaceae. Shrubs were represented by Sapindaceae, Acacia, Myoporaceae and Caesalpinaceae.

The level of Chenopodiaceae/Amaranthaceae (65%) increased in the middle of the midden (Figure 7.8). Eucalyptus decreased and other tree taxa included Melaleuca, Casuarinaceae and Cupressaceae. The levels of Cyperaceae (1%) and Poaceae (2%) were lower while highest levels in the herbaceous taxa included 6% Cruciferae and 3% Onagraceae.

Levels of Chenopodiaceae/Amaranthaceae (52%) were lower in the top of BR3 and the signal from Eucalyptus (6%) was similar to levels in the base section. There was not much difference in the levels of shrub taxa throughout the entire midden. Asteraceae (10%) and Apiaceae (4%) increased in this section while levels of other herbaceous taxa comprising Malvaceae, Convolvulaceae, Cruciferae, Liliaceae, Onagraceae and Haloragaceae were low. Poaceae (3%) and Cyperaceae (3%) returned to levels that were recorded in the base section (Figure 7.8).

Leporillus apicalis faecal pellets were present in the macrofossil assemblage from BR3 TOP. Bones included the broken head of a femur from a species of rodent, and the lower left Ml molar of Pseudomys gouldii (medium sized mouse). There were leaf fragments of Sida petrophila (Malvaceae), Ptilotus obovatus (Amaranthaceae) and Dodonaea viscosa (Sapindaceae). Casuarina cristata cladodes were abundant in the midden matrix (although low levels of this pollen) and there was a selection of seeds including Triodia scariosa ssp. bunicola and Eriochloa sp. Unidentified insect fragments, Coleoptera elytra and Diptera pupae cases were present.

The macrofossil assemblage from the middle section of BR3 was less diverse than the top (Table 7.14). There were Leporillus spp. faecal pellets in the midden and a few strands of cream coloured hair. A right upper incisor (most likely Leporillus apicalis) was identified on the basis of the width, thickness, shape and wear facet on the sub fossil material (Medlin pers.comm 1995). Plant macrofossils included Sida petrophila (Malvaceae), Ptilotus obovatus (Amaranthaceae), Dodonaea sp. and Acacia sp. leaf -214-

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fragments. Casuarina cristata cladodes were present in this section of midden as well as Triodia scariosa ssp. bunicola seeds and Daucus glochidiatus fruits. Insect fragments consisted of Coleoptera sp. appendages.

BR3 BASE macrofossils included juvenile and adult Leporillus spp. faecal pellets. Sida petrophila (Malvaceae) and Acacia sp. leaf fragments were present and a mix of unidentifiable leaf fragments. Casuarina cristata cladodes were the other component in the botanical remains.

7.3.4 Brachina Gorge 4 (BR4) Midden

BR4 Leporillus midden was sub sampled at 10 cm intervals from top to base, and macrofossils from each sample were AMS dated (Table 7.12). Ages ranged from approximately 5 700 years BP. to 7 700 years BP., with age differences between faecal pellets and leaves from the top and base sections. The dates for the 20cm through to 50cm samples appear to be in stratigraphic order.

There was 32% Chenopodiaceae/Amaranthaceae pollen at 60cm. Levels of herbaceous taxa included 13% Asteraceae, 6% Apiaceae, 4% Thymelaceae and lower levels of Convolvulaceae, Liliaceae and Cruciferae (Figure 7.9). Shrub pollen included 6% of Solanaceae, 5% of both Myoporaceae and Fabaceae and 4% of both Sapindaceae and Caesalpinaceae. Highest recorded levels of tree taxa included 4% Eucalyptus and 3% Cupressaceae. There was no record of Cyperaceae at this depth and 2% of Poaceae. One percent Boraginaceae pollen is an indication of either contamination of the midden or a result of excavating samples in the field.

There were lower levels of Chenopodiaceae/Amaranthaceae (26%) at 50cm. The signal from tree pollen had increased, with 9% Eucalyptus and 4% of both Casuarinaceae and Cupressaceae. Levels of shrub taxa also increased with 9% Fabaceae, 8% Myoporaceae, 4% Sapindaceae and 1% Caesalpinaceae, Acacia and Goodeniaceae. There was 10% of Asteraceae and other herbaceous taxa included Thymelaceae (5%) and Apiaceae (3%). Levels of Cyperaceae (6%) and Poaceae (7%) had increased.

The highest level of Chenopodiaceae/Amaranthaceae (58%) was recorded at 40cm while tree taxa was lowest comprising 3% Eucalyptus, and weaker signals of Cupressaceae, Casuarinaceae and Melaleuca. Levels of shrub taxa also decreased, with 4% Myoporaceae, Fabaceae and Solanaceae. There was a decrease in Asteraceae (3%), while Apiaceae returned to levels similar to the record at 60cm and other herbaceous taxa were less than 1% of the pollen sum. Levels of Cyperaceae (3%) and Poaceae (3%) had decreased at this depth (Figure 7.9). -216-

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At a depth of 30cm there was 4% Casuarinaceae and the level of Eucalyptus (3%) was unchanged. There was a marked decrease in Chenopodiaceae/Amaranthaceae (25%). The level of Sapindaceae (14%) increased at this depth while other shrub taxa were present at lower levels and included Acacia, Myoporaceae, Euphorbiaceae, Caesalpinaceae, Solanaceae and Leptospermum (Figure 7.9). There was an increase in Apiaceae (15%) at this depth, while Asteraceae (4%) remained similar to levels recorded at 40cm. Signals of Poaceae (8%) and Cyperaceae (6%) both increased.

There was an increase in the signal from Chenopodiaceae/Amaranthaceae (41%) at 20cm and levels of tree taxa increased (although not to the level at 50cm), with 7% Eucalyptus and 3% Cupressaceae. There was an increase in the level of Myoporaceae (6%), and Acacia (4%) at this depth (Figure 7.9). Apiaceae (3%) decreased while Asteraceae (6%) and Thymelaceae (5%) remained similar to signals at 30cm, and Liliaceae (5%) increased. Poaceae (2%) and Cyperaceae (3%) levels decreased in this section of the midden.

There was 40% Chenopodiaceae/Amaranthaceae pollen at a depth of 10cm. Eucalyptus (6%) was similar to previous levels while other tree taxa were less abundant (Figure 7.9). Levels of shrub pollen had decreased, with the most abundant signals from Fabaceae (7%) and Solanaceae (5%). Apiaceae (7%) increased at the top of the midden and Asteraceae (8%) and Thymelaceae (5%) remained similar to levels at 20cm.

Leporillus spp. faecal pellets, a mix of leaf fragments from the Malvaceae, Dodonaea viscosa ssp. angustrissima (Sapindaceae) leaves, Sida petrophila (Malvaceae) leaves and Casuarina cristata (Casuarinaceae) cladodes were recovered from all depths. Daucus glochidiatus (Apiaceae) fruits and Ptilotus obovatus (Amaranthaceae) leaves were present from 10cm through to 30cm and there were insect fragments in the 10cm, 30cm, 40cm and 60cm samples (Table 7.14). Taxa represented by plant macrofossils were also recorded in midden pollen. However, shrub taxa such as Fabaceae and Solanaceae that were dominant in the pollen, were absent from macrofossil assemblages. Bones from a depth of 10cm included unidentifiable fragments and a lower left incisor from a species of rodent (similar in size to Pseudomys bolami/hermannsburgensis). A right upper incisor from a mouse similar to Pseudomys gouldii was recovered from the 50cm sample.

7.4 Conclusion

A variety of palaeoecological records have been presented in this and the previous chapter, which provides the basis for reconstructing Holocene environments for sites in the northern and central Flinders Ranges. The following chapter is a discussion of the -219-

methodological issues apparent in midden analysis, that is a result of the taphonomy of middens and the nature of different palaeoecological records derived from these deposits. Holocene vegetation communities and palaeoclimates are then discussed at local (site specific) and regional scales and within the context of other palaeoecological research within Australia. -220-

Chapter 8: Methodological Issues and Interpretation of Holocene Vegetation

8.1 Introduction

This discussion chapter addresses the two themes of this research: methodological issues in midden analysis and the contribution of midden macrofossil and pollen records to knowledge of Holocene vegetation communities in the semi-arid zone. The first section of the chapter deals with three main methodological issues in stick-nest rat midden studies. This is followed by a discussion of spatial and temporal trends in palaeoecological records from the northern and central Flinders Ranges by comparing midden pollen spectra with vegetation and midden pollen with macrofossil assemblages. I use the terms Early (7 000- 5 000 B.P), Middle (4 000-2 000 B.P) and Late Holocene (1 000 B.P to present) in the very specific context of the stick-nest rat midden samples throughout this and the following chapter.

8.2 Methodological Issues

Variability in the midden palaeoecological records may of course be influenced by processes other than vegetation change over time. Of particular concern are changing patterns of pollen representation with changes in source areas, and the influences of rat activity on pollen and macrofossil representation. This chapter endeavours to tease apart these different processes in order to interpret changes in vegetation across three time slices of the Holocene.

The chronology of middens is investigated using Accelerator Mass Spectrometry (AMS) Radiocarbon dating of midden macrofossils and pollen. This provides some insight into the taphonomy of different types of deposit, and the possible influences of the rats. The next section compares the fossil pollen record from middens with the modern vegetation and modern cave pollen. This analysis raises questions which are investigated further in section 8.2.3, where fossil pollen and macrofossils are compared. I argue that, with certain caveats, it is possible to separate the sources of variability and make broad inferences about temporal vegetation change, which I do in section 8.3 and 8.4.

8.2.1 Accelerator Mass Spectrometry (AMS) Radiocarbon Chronology

Three middens from Brachina Gorge have been used as case studies to investigate the chronology of midden deposits. AMS dating of leaves, faecal pellets and pollen concentrates from sub-samples of middens were compared to determine the timing of accumulation and chronology of deposits. Processing of middens to release leaves and -221-

faecal pellets is outlined in section 4.9.1 and extraction of pollen concentrates for AMS dating in section 4.11.1 in Chapter 4. Middens BR1, BR3 and BR4 were chosen as they represent different midden types according to the classification scheme in Head et al. (1997) and as a result, can add to the overall assessment of the palaeoecological significance of middens. BR1 and BR3 are examples of amberat middens and BR4 is a free standing midden with apparent layering throughout the deposit. The AMS dates for each midden have been summarised in a series of scatter plots (Figure 8.1 and Figure 8.2).

In BR3, a Mid-Holocene midden, the age of leaves (3 570 ± 120 BP OZB290) from the top sample were contemporary with faecal pellets (3 790 ± 90 BP OZB300) within the reported sigma errors. Ages of leaves and faecal pellets from the base sample (3 690 ±180 BP OZB298 and 3 640 ± 70 BP OZB293 respectively) were also comparable (Figure 8.1). Within the context of the entire deposit (a depth of 60cm), it is apparent that the deposition of this midden was rapid.

Leaves and faecal pellets from top and base samples of the Early Holocene midden (BR1) were dated at different laboratories. Ages of faecal pellets from the top and base (7 400 ± 90 BP BETA8108 and 7 310 ± 90 BP BETA8109 respectively) are contemporary within the 1 sigma errors but are older than leaves from the same samples (6 800 ± 70 BP OZB292 and 6 710 ± 80 BP OZB291 respectively) dated at a different laboratory. Ages of faecal pellets suggest that this midden deposit accumulated rapidly, as was the case for the Middle Holocene (BR3) midden (Figure 8.1). Difference in the ages between faecal pellets and leaves from the same samples may result from reworking of this material through the deposit during construction and occupation of the midden by the rat. It is also a possibility that differences in ages are attributed to systematic discrepancies between laboratories.

BR4 midden displayed apparent layering from the top to the base of the deposit, over a depth of 60cm. Radiocarbon dates were obtained from two different laboratories. Faecal pellets and leaves from the top of the midden were contemporary within reported errors (6 930 ± 80 BP BETA81091 and 6 290 ± 190 BP OZB295 respectively) while another date on faecal pellets from ANSTO (5 690 ± 70 BP OZB294) was younger (Figure 8.2). Changes in the depositional history of the midden have been detected by the series of AMS dates on leaf macrofossils from successive layers. Ages of leaves from the 50cm through to 20cm layer (distance from the base towards the top of the midden), are in stratigraphic order (Figure 8.2). Ages of the 60cm, 50cm and 40cm layers are contemporary within the reported 1 sigma errors and therefore indicate a period of rapid accumulation. Following this there is a difference of approximately 300 years between the 40cm and 30 cm layer of midden which dates at 6 580 ± 60 BP (BETA 87300) (Figure -222-

7500- • (BETA8108) _p X • 0. 7250- X cd faecal pellet in (BETA8109) u 7000- >* leaf U T (OZB292) SO A 6750- T < _L A X 6500- 1 (OZB291) 1

Midden sub-samples

(OZB300) 3800- • (c)ZB298 )

CD m faecal pellet in 3700- 1 ] CO 1[ u 3600- T A leaf u on A (OZB 293) < 3500- (OZB2991) 3400- 1 1 4> 1 / < »* Midden sub-samples

Figure 8.1: Scatter plots of AMS dates on faecal pellets and leaf macrofossils from top and base samples of Brachina Gorge 1 and Brachina Gorge 3 middens. -223-

TOP 6 930 ± 80 yrs B.P 5 690 ± 70 6 290 ± 190 2 890 ± 60

6 520 ± 50 6 320 ± 60

layering 6 580 ± 60 6 600 ± 60

Unconsolidated 6 880 ± 50 sediment 6 940 ±60 Compacted sediment Leporillus sp. 6 920 ± 50 faecal pellet 6 150 ± 50

6 610 ± 90 Twigs and leaf 7 720 ± 90 macrofossils 6 650 ± 80 ASE7 060±5()

ROCK LEDGE

cm A. AMS date on leaf macrofossils • AMS date on pollen i^p limestone

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tU 5000- u O pollen < 4000- 3000- O

2000 —' 1 i 1 l 1 i ^ # # f 4>

Midden Depth

Figure 8.2: Brachina Gorge 4 midden displays apparent layering. AMS dates on faecal pellets from the top and base samples and AMS dates on leaf macrofossils and pollen concentrates at 10cm intervals are illustrated on the schematic diagram and scatter plot. -224-

8.2). It is possible that the midden was abandoned for a period of time and then re- occupied. Age of faecal pellets and leaves from the base sample (6 610 ± 90 BP OZB296 and 6 650 ± 80 BP OZB297 respectively) were contemporary within reported errors while the BETA81092 date on faecal pellets was older (7 720 ± 90 BP BETA81092). On the basis of the 7 720 ± 90 BP age for the base and 6 290 ± 190 BP for the top section, BR4 midden has accumulated over a few hundred years. On the whole, different macrofossils (leaves and faecal pellets) from the same sample of midden were found to be of a similar age. However, contamination by younger or older material is always a possibility that must be recognised.

AMS dates on pollen concentrates from BR4 again reveal the complexity in the temporal resolution of midden deposits, as AMS dates for the 10cm and 50cm layers are problematic (Figure 8.2). Given that the macrofossil dates are in stratigraphic order, the 10 cm layer pollen date of 2 890 ± 60 BP (BETA92926) can be attributed to possible reworking of younger pollen due to remobilisation of amberat in more humid periods. Alternatively, contamination could have occurred during processing of the sample in the laboratory. The 50cm layer pollen date of 6 150 ± 50 BP (BETA92927) may be an indication of reworking of material when the midden was occupied. The rats themselves may have moved through the nest/midden, mixing pollen rich sediment through the layered midden (Head et al 1997). The ages of pollen concentrates and leaf macrofossils from each layer of midden are not always contemporary. In the 20cm and 60cm layers, pollen is younger than macrofossils and in the 30cm and 40cm layers pollen and macrofossils are contemporary (Figure 8.2).

The case studies from Brachina Gorge indicate that refining the temporal resolution of stick-nest rat middens remains problematic. Generally, there is still uncertainty over contamination and reworking of material. This complexity in our understanding of the temporal resolution of middens illustrates some of the problems encountered when working with these deposits. That is, middens can be stratigraphically complex and there is the possibility of breaks in the depositional history or collapse in the structure as the deposit is constructed (Elias 1990; Spaulding etal. 1990; Webb and Betancourt 1990).

AMS radiocarbon dating of leaves, faecal pellets and pollen concentrates suggests that accumulation of middens can be a rapid event, however time of accumulation does vary from site to site. The deepest deposits (approximately 60cm depth) have formed within a few hundred years. AMS dates on samples from top, middle and base sections of middens have (in most cases) indicated that different macrofossils (leaves and faecal pellets) returned contemporary dates within the reported 1 sigma errors. Where discrepancies occurred, this has been attributed to different laboratory results or reworking of younger or older material throughout the midden. -225-

As a result of the investigation into taphonomy of middens from the Brachina Gorge case studies, the following factors are important to consider, when using midden pollen and macrofossil records for palaeoenvironmental reconstructions:

1. Generally, different types of macrofossils from the same sample of midden are a similar age, thus providing a reliable age for a midden. 2. The accumulation of a midden is a rapid event. Given the different types of midden (eg. amberat midden, layered midden, stick-nest described in Head et al. 1997) and variability in the amount of 'dateable' material available, single dates from a midden should provide a 'ball-park figure' age for the deposit. 3. Further investigation is required into the contemporaneity between pollen and macrofossils in middens. This study indicates that pollen is contemporary with associated macrofossils in some parts of a midden, while in other layers there is a difference of a few hundred years (excluding examples of contamination by younger pollen). The reliability of pollen records from middens can be justified in the context of the method used for extracting pollen, when processing the midden (section 4.91. in Chapter 4). The pollen represents an averaged record for the relevant sub-sample of midden because of the way that it was extracted from the residue soak, and therefore, the dates are imprecise but reliable.

Therefore within the context of my study, each midden can be regarded as representing a defined time-slice within the Holocene, providing specific palaeoecological information at a number of different locations in the central and northern Flinders Ranges. Records from individual midden sites can be synthesised into a regional reconstruction of palaeovegetation and climates during the Holocene. While there might be concerns over dating of any individual sample of midden, the thorough spatial and temporal coverage of sites should overcome these.

8.2.2 Comparison Between Midden Pollen and Modern Vegetation

An investigation of what pollen taxa are being recorded in middens relative to source distance in the vegetation is presented in this section. I aim to determine what mix of local and regional pollen is being captured in middens by referring to data on vegetation surveys at midden sites, and findings from the cave modem pollen study in Chapter 5.

Arkaroola-Mount Painter Sanctuary Middens The number of pollen taxa recorded in midden assemblages ranged from 17 to 27 and was consistently higher than the number of taxa recorded in vegetation surveys (Table 8.1). North Well Creek 1 (560 yrs B.P) and Radium Creek 2 (2 480 yrs B.P) midden pollen assemblages were more diverse than the surrounding vegetation. At North Well Creek 1 -226-

there were ten taxa recorded in the vegetation including shrubs and two herbaceous taxa (Table 8.1), while midden pollen included trees (eg. Eucalyptus and Melaleuca) and a more diverse assemblage of shrubs (eg. Leptospermum, Acacia, Myoporaceae and Euphorbiaceae) and herbaceous taxa (eg. Apiaceae, Liliaceae and Convolvulaceae). At Radium Creek 2, Melaleuca, Asteraceae, Malvaceae and Chenopodiaceae were recorded in surveys while midden pollen contained these taxa in addition to other tree, shrub (eg. Acacia .and Caesalpinaceae), herbaceous (eg. Convolvulaceae, Thymelaceae and Liliaceae) and grass taxa (Table 8.1). Some of these taxa are poorly represented in the modem pollen rain at cave sites (see Chapter 5) thus indicating more extensive cover of shrubs and herbs around 2 000 yrs B.P. At both sites, the middens were located in exposed conditions at the top of rocky outcrops, with a sparse vegetation cover on the steep slope outside each overhang. The position of middens at the front of overhangs also provided ideal conditions for the capture of windblown pollen onto midden surfaces.

Other middens were located in protected sites at the rear of caves or along crevices in the side of cave walls. There was similarity between taxa recorded in the midden pollen and taxa present in the modem vegetation. Waterfall 1 site, with an AMS date of post 1950, is an analogue for present day vegetation. Six taxa were recorded in both the midden pollen and vegetation cover at the site (Table 8.1) and there were additional tree (Eucalyptus, Casuarinaceae and Melaleuca) and shrub (Fabaceae and Euphorbiaceae) pollen taxa in the midden assemblage, representing possible extra-local and regional sources of windblown pollen. At North Well Creek 2 (3 670 yrs B.P), taxa recorded in the vegetation surveys (10 taxa representing shrubs, herbs and grasses) were also present in the midden pollen with the exception of Proteaceae and Goodeniaceae (Table 8.1) known to be low pollen producers and poorly dispersed. For Radium Creek 1 (4 350 yrs B.P), nine out of 13 taxa recorded in the vegetation were also present in the midden pollen. Differences resulted from more diverse herbaceous taxa (eg. Liliaceae, Apiaceae, Convolvulaceae and Onagraceae) recorded in the midden and Proteaceae, Solanaceae, Lamiaceae and Cucurbitaceae present in the vegetation but absent from the midden (Table 8.1). Shrub taxa (with the exception of Lamiaceae and Santalaceae) were also recorded in the pollen assemblage at Oppaminda Track (880 yrs B.P), but like other middens, there was a large component of herbaceous taxa recorded in the midden that was absent from the immediate vegetation at the site (Table 8.1). Similar trends were apparent in the records from Haematite Hill 1 (590 yrs B.P). The vegetation was dominated by shrubs (six different taxa) and species of Asteraceae and Malvaceae. However in addition to this, midden pollen contained another twelve shrub and herbaceous taxa including Euphorbiaceae, Goodeniaceae, Lamiaceae, Liliaceae and Haloragaceae (Table 8.1). Ten out of the eleven taxa recorded in the vegetation at Radium Creek 3 (420-450 yrs B.P) site were present in -227-

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the midden pollen and unlike the majority of other sites, herbaceous taxa in the pollen was less diverse. Ten out of twelve taxa from the vegetation at Arkaroola 1 were recorded in the midden pollen, with Proteaceae and Malvaceae exclusive to the vegetation (Table 8.1).

Mount Chambers Gorge and Brachina Gorge Middens Middens from the south-western face of Mount Chambers were located in small caves opening out onto sparsely vegetated scree slopes. The midden pollen contained a larger number of genera representing a mix of local and regional taxa with a maximum of 31 pollen types recorded, compared to a maximum of 14 taxa listed in vegetation surveys.

At Mount Chambers 1 site (410 and 1 280 yrs B.P), species of Acacia, Sapindaceae, Lamiaceae, Malvaceae and Chenopodiaceae/Amaranthaceae were recorded in the vegetation and Acacia was the only one that was not present in the midden pollen (Table 8.2). Very low levels of trees (eg Cupressaceae, Eucalyptus .and Melaleuca), shrubs (eg. Caesalpinaceae and Goodeniaceae) and herbaceous taxa (eg. Thymelaceae and Convolvulaceae) were exclusive to the midden pollen and most likely represented extra- local windblown sources and ephemeral herbs being captured in the midden. Additional herbaceous taxa in the pollen may have included ephemeral species that were absent at the time of survey. At Mount Chambers 2 site (230 and 590 yrs B.P), vegetation cover outside the cave was more diverse with nine taxa including trees, shrubs, herbs, chenopods and grasses recorded. Acacia and Pittosporaceae were not in the midden pollen but shrub and herb taxa that were present in the pollen at Mount Chambers 1 were also recorded at this site (Table 8.2).

At Chambers Gorge MD3 site (7 260 yrs B.P) at the eastern end of the gorge, Sapindaceae and Myoporaceae were present in the vegetation and midden pollen and the presence of an additional seven shrub taxa were exclusive to the midden (Table 8.2). Taxon including Caesalpinaceae, Proteaceae and Solanaceae that are usually under represented in modem pollen rain (Chapter 5), were present in the midden even though not in the modern flora. Six herbaceous taxa were also only recorded in the midden (Table 8.2). There was a more extensive cover of herbs in the vegetation at this time. At Chambers Gorge 1 site (470 yrs B.P) inside the gorge, six out of the seven taxa present in the vegetation were also recorded in the pollen. The midden pollen lacked a diverse herbaceous component unlike other middens from the Mount Chambers Gorge study site (Table 8.2). Shrub taxa recorded in the midden (eg. Caesalpinaceae) was not at the site but abundant at other sites throughout the gorge and thus possibly represents an extra- local source.

At Brachina Gorge 1 (6 700-7 400 yrs B.P), 14 taxa in the vegetation survey, comprising shrubs, herbs, chenopods and grasses were also recorded in the midden pollen (Table -230-

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8.3). Trees were exclusively represented in the pollen as were two additional shrub and five herbaceous taxa. The herbs (eg. Convolvulaceae, Liliaceae) are known to be poorly represented in the modern pollen rain (Chapter 5) but are recorded at higher levels during this part of the Holocene as is the case at other Early Holocene sites. The vegetation at Brachina Gorge 2/7 (5 020-5 790 yrs B.P) was similar to Brachina Gorge 1. Again, trees were not recorded at the midden cave site but Cupressaceae, Casuarinaceae, Eucalyptus and Melaleuca were present in the pollen assemblages (Table 8.3). At Brachina Gorge 3 (3 640-3 690 yrs B.P) and Brachina Gorge 4 (6 290-7 720 yrs B.P), seven shrub taxa recorded in midden pollen were not present in vegetation surveys and there was also a diverse assemblage of herbaceous taxa (eg. Thymelaceae, Convolvulaceae, Onagraceae, Haloragaceae) exclusive to midden pollen.

In summary, there were some consistent differences between the composition of midden pollen and vegetation at midden sites. Midden pollen, especially in the Early Holocene (7 000-5 000 yrs B.P) middens, displayed a more diverse herbaceous assemblage than the vegetation surveys. Tree taxa were also commonly recorded in midden pollen assemblages when absent from the midden site. It has been demonstrated that midden pollen is likely to be recording a mix of local taxa at the cave sites that mirrors what the modern pollen rain studies have shown in Chapter 5, in addition to inclusion of taxa from outside the immediate cave environments from both extra-local and regional sources. It is questionable as to whether taxa represented in middens that are known to be poorly dispersed in the modern pollen rain, are an indication of vegetation change over time or inclusion of semi-arid ephemeral species that may not have been present at the time of the vegetation surveys. Calculation of Macrofossil:Pollen Index values in the next section may help separate out these possibilities.

8.2.3 Comparison Between Fossil Pollen and Macrofossils in Middens

MacrofossihPollen indices (MPI) were calculated for ten taxa that were represented in both pollen and macrofossil records in middens from the Arkaroola-Mount Painter Sanctuary and Brachina Gorge. Mount Chambers Gorge middens have MPIs calculated for four taxa as these middens were not rich in macrofossils (Table 8.4). Calculation of MPI values investigates the relationship between pollen and macrofossil records in middens. MPI values are based on an aggregate measure of middens from each study site. The common taxa are representative of tree, shrub, herb, chenopod and grass vegetation categories. The pollen is scaled (in relative percentage) to approximate the relative abundance scale used for plant macrofossils. The index is calculated by "...summing the differences between the relative abundances of macrofossil and pollen for each taxon from all middens in which that taxon was represented by either data type and dividing by the number of samples..." (Anderson and Van Devender 1991:17). According to Thompson -234-

(1985), the index should vary from -5 (taxa represented at levels greater that 50% in the pollen data and lacking plant macrofossils) to +5 (taxa are abundant in plant macrofossils and absent from the pollen). MPI values between -1.00 and +1.00 indicate pollen and macrofossils are likely to be showing similar vegetation signals (Anderson and Van Devender 1991). For calculations of MPI values, see Appendix 9. In middens, macrofossils were ranked as 1 (rare) or 2 (uncommon) on a relative abundance scale, in contrast to pollen that was ranked anywhere from 1 (rare) to 5 (abundant) (Appendix 9).

All MPIs were negative, indicating that taxa were better represented by pollen rather than macrofossils. Index values ranged from -3.8 (Chenopodiaceae/ Amaranthaceae) to -0.5 (Malvaceae) and these maximum and minimum values occurred in Mount Chambers Gorge middens. MPI values for each taxon were assigned to a vegetation category and discussed below.

Common Taxa Arkaroola Sanctuary Mount Chambers Gorge Brachina Gorge Casuarinaceae -1.15 * -1.44 Acacia -0.7 * -1.06 Myoporaceae -1.61 -1.83 -1.88 Sapindaceae -1.54 * -1.25 Caesalpinaceae -1.62 * * Apiaceae -1.77 * -1.56 Asteraceae -2.7 * -2 Malvaceae -0.85 -0.5 -0.56 Chenopodiaceae -2.3 -3.8 -3.69 Poaceae -1.61 -1 -1.88 macrofossil taxa not recorded in any midden from the study area.

Table 8.4: Macrofossil:Pollen Indices for middens from the Arkaroola-Mount Painter Sanctuary, Mount Chambers Gorge and Brachina Gorge.

Trees MPI values for Casuarinaceae (-1.15 and -1.44) were similar at Arkaroola and Brachina Gorge (Table 8.4). Pollen values ranged from <1% - 2% for middens at these sites, and macrofossils (cladodes) were more common in the suite of middens from Brachina Gorge, compared to the middens from Arkaroola. -235-

Shrubs Arkaroola middens recorded a more similar representation of Acacia pollen and macrofossils (MPI -0.7) in the middens, compared to Brachina Gorge middens. Levels of this pollen were low ( <1% -2%) in most middens and macrofossils (leaves and pods), when present, were also recorded in low abundances. Levels of Myoporaceae pollen were variable, ranging from <1% to 15%. Macrofossils (leaves) were less common in midden samples (one Mount Chambers Gorge midden and three middens from Brachina Gorge and Arkaroola), resulting in MPI's ranging from -1.61 to -1.88 (Table 8.4). Representation of Sapindaceae in pollen and macrofossils was more similar at Brachina Gorge (MPI -1.25) compared to Arkaroola (MPI -1.54). This is another taxon where the representation of pollen was variable in middens, ranging from <1% in Mount Chambers Gorge middens to 15% in some of the middens in the Arkaroola-Mount Painter Sanctuary. Caesalpinaceae macrofossils (stem and seed pods) were only recorded in two middens from the Arkaroola-Mount Painter Sanctuary (MPI -1.62).

Herbs MPI values of -2.7 and -2 were calculated for Asteraceae from Arkaroola and Brachina Gorge middens respectively, indicating a stronger representation of this taxon in midden pollen (values having been recorded as high as 49%). Cassinia laevis leaves and flowers were found as macrofossils and ranked 1 (rare) on the relative abundance scale. Macrofossils of the genus Apiaceae, were Daucus glochidiatus seeds and fruits, and found in Arkaroola and Brachina Gorge middens. The representation of this taxon in midden pollen was variable with some middens recording <1% and others up to 21%. This resulted in MPI values of -1.56 and -1.77 (Table 8.4). A smaller MPI value for Malvaceae (-0.5 to -0.85) from middens at all study sites indicated a more comparable representation of macrofossils and pollen in middens. Pollen levels were low (1%), given that Malvaceae is known to be a poorly dispersed and relatively large pollen grain. Leaf fragments of Malvaceae were common building material used by the stick-nest rat and recorded in macrofossil assemblages of all middens.

Chenopods MPI values for Chenopodiaceae/Amaranthaceae (MPI -3.8 to -2.3) indicate that this taxon dominated midden pollen assemblages and was less abundant as a macrofossil (Table 8.4). There were higher levels of this pollen in the Mount Chambers Gorge and Brachina Gorge middens from the central ranges. Chenopodiaceae/Amaranthaceae pollen is known to be produced in large amounts and regionally dispersed, in addition to the fact that the taxon is commonly found in vegetation communities at midden cave sites. Extreme levels of this pollen (especially in Mount Chambers midden samples), must be generated from a -236-

combination of the local vegetation within the foraging range of Leporillus and a windblown regional component.

Grasses Poaceae MPI values were similar at each study site (Table 8.4). Midden pollen signals ranged from l%-8% and macrofossils included small numbers of seeds ranking 1 on the relative abundance scale.

Middens from all three study sites were grouped into Early (7-5ka), Middle (4-2ka) and Late Holocene (

Common Taxa (5-7ka) Middens (2-4ka) Middens (

Table 8.5: MacrofossihPollen Index for Early, Middle and Late Holocene middens. -237-

Pollen assemblages are more abundant and diverse than macrofossils in middens as evidenced by MPI values. Therefore, macrofossils can be regarded as a secondary source of information available in Leporillus spp. middens. They can confirm local occurrences of species where it is not possible to distinguish a local or regional source for the pollen taxon. There is some variability in MPI values from Arkaroola, Mount Chambers Gorge and Brachina Gorge as levels of pollen taxa and macrofossil type and abundance were to some extent unpredictable.

For key selected taxa, a table summarising the representation of each taxon in relation to levels recorded in the modern pollen rain, midden pollen spectra, and presence of macrofossils is presented (Table 8.6). These features are important for the interpretation of vegetation histories from the sites.

Key Taxa Variability in modern pollen rain, midden spectra and macrofossils Casuarinaceae • low to moderate levels in cave pollen traps irrespective of whether present or absent in the vegetation • levels in midden pollen similar to modem pollen spectra in cave traps • macrofossils confirm local presence of taxa at sites Acacia • poorly represented in the modem pollen spectra at cave sites • levels in midden pollen similar to modem pollen spectra in cave traps • macrofossils similar signal to midden pollen ! Myoporaceae • low levels in modem pollen spectra at cave sites • variable levels in midden pollen spectra when present in the vegetation at the site • lower levels of macrofossils compared to midden pollen Sapindaceae • variable levels in cave pollen traps when vegetation is present • range of levels in midden pollen are similar to the modem pollen rain • macrofossils similar signal to midden pollen spectra Caesalpinaceae • variable levels in the modem pollen rain from cave trap sites • recorded in the midden pollen spectra when present in the vegetation cover • macrofossils not common in middens -238-

Apiaceae • high levels in the modem pollen rain when not necessarily present in the vegetation • similar levels in the midden pollen spectra when not present at the site • presence of macrofossils in middens Asteraceae • can be over represented in the modem pollen rain • variable levels in the midden pollen spectra irrespective of whether present or absent in the vegetation • higher levels of pollen compared to less abundant macrofossils Malvaceae • poorly represented in modem pollen rain at cave sites • low to moderate levels in midden pollen spectra • commonly found as macrofossils in middens Chenopodiaceae • variable levels in modem pollen rain • higher levels in midden pollen spectra • macrofossils less common in middens Poaceae • well represented in modem pollen at cave sites • variable levels in midden pollen • rare occurrence of macrofossils in middens

Table 8.6: Summary of representation of key taxa in the modem pollen rain, midden pollen and macrofossils.

8.3 Regional Patterns in Pollen Data for Individual Taxa

Holocene vegetation communities can be investigated by discussing regional patterns of individual pollen taxa in the central and northern ranges. Average values for key taxa in middens were mapped for three periods in the Holocene, lka to the present (12 middens), 4-2ka (6 middens) and 7-5ka (8 middens). The taxa included representatives from tree, shrub, herb, grass and chenopod vegetation categories. Levels of these taxa in the modem pollen were included, as a modem equivalent for comparison with the fossil records.

Cupressaceae was consistently represented at low levels in all pollen records in the Holocene (Figure 8.3). There were similar levels recorded in the Lake Frome study (Singh and Luly 1991) and other middens at Arkaroola and Mount Chambers Gorge from a previous study (McCarthy et al. 1996). The levels of this taxon in the modem pollen from the northern ranges were more variable than levels of the fossil pollen. Levels in the modem and fossil records were more similar in the central ranges (Figure 8.3). Cupressaceae macrofossils were rare in middens from all study sites. -239-

Levels of Casuarinaceae show no variation throughout the Holocene ranging from <1-1% (Figure 8.3). The record from Lake Frome was more abundant during the Early to Mid- Holocene period while the modem equivalent was comparable to the midden records. Cladode macrofossils confirmed the local occurrence of these trees at the midden sites indicating the pollen signals were not exclusively from a regional source.

Eucalyptus was most abundant in Brachina Gorge at 7-5ka compared to levels at 4-2ka. The Mount Chambers record at 7-5ka, was similar to that at Brachina Gorge. During the 4-2ka period, Eucalyptus levels were higher in the northern ranges, compared to Brachina Gorge in the central ranges. Levels in middens dating lka to the present, were variable and generally higher in the north (Figure 8.3), reflecting riverine woodland communities along creeks and within the confines of gorges. The Lake Frome core recorded higher levels of this taxon for the Late Holocene but there were similar levels in midden and lake assemblages for the Middle Holocene (Figure 8.3). The modem pollen signals were varied as levels of this taxon from riverine woodland sites were higher than records from chenopod shrubland sites. Eucalyptus macrofossils were rare in midden records. The pollen of Eucalyptus is known to be regionally dispersed and levels decline in drier conditions (D'Costa and Kershaw 1997). The decline in Eucalyptus in the fossil record from Brachina Gorge at 4-2ka is a response to the increase in aridity towards the Late Holocene.

Levels of Sapindaceae decreased from the Early (7-5ka) to Middle (4-2ka) Holocene period at Arkaroola, while Middle Holocene (4-2ka) levels at Brachina Gorge remained similar to the Early Holocene (7-5ka) records. Levels from Mount Chambers Gorge in the Early Holocene (7-5ka) were lower than Arkaroola and continued to decline into the Late Holocene period. Sapindaceae levels were variable at Arkaroola (lka to present) and there were lower levels at Brachina Gorge and Mount Chambers Gorge, compared to the Lake Frome record and levels recorded in the modem pollen (Figure 8.3).

Levels of Asteraceae at Brachina Gorge during 7-5ka were higher compared to Mount Chambers Gorge and Arkaroola. There was a drop in levels from the Middle (4-2ka) to Late Holocene (lka to present) at Brachina Gorge, however levels were more abundant in the northern ranges and in the Lake Frome record. There was variability in signals from this taxon at Arkaroola in the Late Holocene, but generally there were higher levels in the northern ranges compared to Mount Chambers Gorge (Figure 8.3).

There were less abundant levels of Chenopodiaceae/Amaranthaceae in the Early Holocene (7-5ka) at Brachina Gorge compared to records from Mount Chambers Gorge. During the Middle Holocene (4-2ka), there were higher levels of this taxon at Brachina Gorge, compared to levels from the northern ranges (Figure 8.3). At 4-2ka, levels at Arkaroola -240-

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were variable but generally less abundant compared to Lake Frome and Brachina Gorge. Mount Chambers Gorge recorded extremely high levels in the Late Holocene that were more abundant and widespread than levels in the middens from the northern ranges and the Lake Frome pollen core (Figure 8.3).

Cyperaceae plus Poaceae signals were combined to represent grassland in the fossil record. Levels at Brachina Gorge were similar from 7-5ka and 4-2ka and there were lower levels at Mount Chambers Gorge at this time. Arkaroola middens at 4-2ka recorded some higher levels compared to sites in the central ranges. Levels in Late Holocene (lka to present) middens from Arkaroola were more abundant compared to records at Mount Chambers Gorge. In previous work at Mount Chambers Gorge, Middle Holocene signals were higher than Late Holocene records. Poaceae was more abundant in the Lake Frome record during 7-5ka and 4-2ka compared to midden pollen records, and decreased in Late Holocene sediments from the core to be similar to the midden records for lka to the present (Figure 8.3).

8.4 Overview of Holocene Vegetation Communities

At 7-5ka, palaeoecological records from Brachina and Mount Chambers Gorge suggest higher levels of trees and shrubs with an understorey dominated by chenopods and herbs (Figure 8.4). A low signal from grasses was consistent across the central ranges at this time. Shrubs were more diverse at Mount Chambers Gorge and there were more abundant chenopods and lower levels of herbs at this site, compared to the same time period at Brachina Gorge. Modern pollen from Brachina Gorge shows lower levels of tree and herb taxa compared to the fossil records (Figure 8.5). There were higher levels of chenopods in the middens while grasses were similar at 7-5ka and the present day. At Mount Chambers Gorge, shrubs were more abundant and widespread in the fossil record compared to modern pollen, while chenopods were lower in the modern pollen, compared to the midden (Figure 8.5). The 7-5ka middens display spatial homogeneities in the vegetation records with similar compositions of trees, shrubs and grasses.

From 4-2ka, chenopods with an herbaceous understorey were dominant in the central ranges (Figure 8.4). Similar abundances of trees and shrubs occurred in the northern ranges at this time. There were lower levels of chenopods and herbs in the modern pollen compared to the fossil record at Arkaroola and similar signals of trees and shrubs (Figure 8.5). At Brachina Gorge, levels of herbs were higher in the fossil record while chenopods were similar to the modern pollen signal. Trees were slightly higher in the modern signal compared to midden records (Figure 8.5). There are spatial homogeneities in samples from the northern ranges, but not between the north and central ranges with respect to the composition of trees and herbs at this stage of the Holocene. Records of -248- -249-

1. 5km west of Haematite Hill: 1995 11. Chambers Gorge camp: 1995 2. 5km west of Haematite Hill: 1996 12. Chambers Gorge camp: 1996

3. Junction of Haematite Hill and North Well Creek 1995 13. Chambers Gorge: 1996 4. Junction of Haematite Hill and North Well Creek 1996 14. Eastern fenceline Chambers Gorge: 1995 5. Nooldoonooldoona Water Hole: 1995 15. Eastern fenceline Chambers Gorge: 1996

6. Echo Camp Water Hole: 1996 16. Western boundary Brachina Gorge: 19% 7. Arkaroola Rd: 1995 17. Opposite BR4 midden site: 1996

8. 5km west of Mount Chambers: 1995 18. Opposite BR1 midden site: 1996

9. 5km west of Mount Chamfers: 1996 19. Eastern end Brachina Gorge: 1995 10. Footslopes of Mount Clmmbers: 1996 20. Eastern end Brachina Gorge: 1996 21. Elatina/Nucdeena Formation: 1995 Figure 8.5: Summary of modern pollen rain data199 for5 an d 1996 from trap sites alona west-east transects across the Flinders Rangesa t Arkaroola, Mount Chambers Gorge and Brachina Gorge -250-

shrubs were more abundant in the northern rather than central ranges while, chenopods were more abundant in the central ranges.

At lka to the present, records from the middens reflected the high degree of spatial variability observed within present semi-arid vegetation communities especially in the northern ranges (see Chapter 3). Variable levels of tree and shrub taxa with an understorey dominated by herbaceous taxa and scattered chenopods largely persisted in the northern ranges. In the central ranges, chenopods were dominant at sites on Mount Chambers, while the midden record from inside the gorge recorded higher levels of trees that is indicative of a riverine woodland community (Figure 8.4). The modern pollen signals at Arkaroola were similar to fossil records. Mount Chambers Gorge fossil records of chenopods showed higher levels in middens compared to the modern equivalent (Figure 8.5) while there were higher levels of tree, shrub and herb pollen taxa in the modern pollen. This variability may be recent, given the homogeneities evident in the earlier midden records. The increased variability in vegetation from the central and northern ranges started some time after the 4-2ka time slice and increased later in the most recent phase from lka to the present.

8.5 Conclusion

Leporillus spp. middens are well dated deposits, providing specific palaeoecological information for a defined period of the Holocene at a number of locations in the central and northern Flinders Ranges. The temporal resolution of middens can be complex however it has been demonstrated that multiple dating of different components can provide a reliable age for the deposit. Pollen taxa recruited into midden assemblages are a mix of local taxa from cave sites and regional taxa outside the immediate environment. MacrofossihPollen Index values indicated that taxa common in both pollen and macrofossil assemblages were better represented by pollen. Macrofossils are a secondary line of evidence that can verify the local occurrence of a specific taxon whose source (either regional or local) may not be able to be distinguished in only the pollen record. Records from individual midden sites can be synthesised into a regional reconstruction of palaeovegetation. Causes for changes and variability in the Holocene vegetation record will be considered in the next chapter. -251-

Chapter 9: Long-term Vegetation Change and Holocene Palaeoclimates from Stick-nest Rat Midden Records

9.1 Long Term Change in Flinders Ranges Vegetation

Long term change in Holocene vegetation communities in the central and northern Flinders Ranges was identified in the previous chapter, and this sets the context for examining possible causes for these changes. This chapter briefly recaps vegetation histories inferred from Leporillus spp. middens and then explores the influence of climate, fire and biogeographical parameters as causes for vegetation change. An interpretation of Holocene palaeoclimates inferred from midden vegetation records concludes this chapter.

Woodland and shrubland communities with herbaceous and grassy understories were dominant at the end of the Early Holocene (7 000-5 000 B.P) in the northern ranges, and shrublands with an understorey of chenopods and herbs were dominant in the central ranges. Shrubland communities declined in the central ranges while persisting in the north from 4 000-2 000 B.P, to be replaced by chenopod shrublands with less abundant herbaceous understories. Chenopod shrublands continued to increase into the Late Holocene (1 000 B.P to the present) in the central ranges. When chenopod shrublands increased markedly in the central ranges (4 000-2 000 B.P), herbaceous taxa (dominantly Asteraceae) decreased. However in the northern ranges an increase in chenopod shrubs was less dramatic, and woodland and shrubland communities were still dominant.

9.2 Causes of Vegetation Change: climate, fire and biogeographical parameters

Semi-arid vegetation communities are determined by climate, fire, moisture availability, soil, and topography, resulting in a high level of spatial variability that produces a mosaic of vegetation communities within the region (a theme explored in detail in Chapter 3). Climate is a significant factor, and because it affects the other parameters, it is discussed first as a cause for vegetation change.

Climate Holocene vegetation records inferred from the Leporillus spp. middens are reconciled (or not) with evidence used to reconstruct Holocene climate change. Elements of climate that are relevant to this discussion (total precipitation, summer/winter rainfall, and variability (ENSO)) are addressed. -252-

The Early Holocene was a complex period during which temperature and precipitation approached modern conditions (Ross et al. 1992). Most rapid change in temperature and rainfall occurred between 12 000 and 9 000 years B.P, with an expansion of woodland vegetation in southern Australia. This was followed by another peak in effective precipitation between 8 000 and 4 000 years B.P with a further increase in woodland and forests. The 6 000 years B.P time slice is well marked in most records of Holocene environmental change (Wasson et al. 1991). Temperature in Australia was higher than present and lake levels were highest since the pre-last glacial maximum lacustral phase (Ross et al. 1992). Climate was warmer than the present and rainfall increased in the north while decreasing in the south of the continent, thus trending towards high summer and low winter rainfall (De Deckker et al. 1988). The bulk of evidence from the Late Holocene (3 000 years BP), suggests cooler and drier conditions (Ross et al. 1992). Overall, climatic variability has been a major feature of the Late Holocene (Kershaw 1995).

Total Precipitation and Temperature Change Lower temperatures and reduced levels of precipitation are characterised by steppe and heath vegetation communities and higher levels than the present of Chenopodiaceae (Kershaw 1995). During periods of warmer temperatures and effectively wetter conditions, there is an expansion of woodland and forest communities dominated by Eucalyptus and Casuarinaceae (Crowley and Kershaw 1994; Lloyd and Kershaw 1997). Present day environments in the Flinders Ranges indicate widespread aridity and climatic variability with vegetation communities dominated by sclerophyllous woodlands, shrublands and chenopod shrublands. The response of vegetation to changes in total precipitation is reflected in the midden records. There were woodland and shrubland communities present at the end of the Early (7 000-5 000 B.P) and Middle Holocene (4 000-2 000 B.P) (indicative of wetter conditions), to be replaced by chenopod shrublands in the Middle to Late Holocene (4 000 B.P to the present) in the central ranges and shrublands and a less extensive cover of woodlands in the northern ranges. A Middle-Holocene (6 000-5 000 B.P) arid phase (theme explored in Chapter 2) is not supported by the midden records.

Seasonality: Summer/Winter Rainfall According to Kershaw (1995), more data is needed to further test the proposal of a shift from winter to summer rainfall across central Australia in the Early Pleistocene or Late Holocene as suggested from the Lake Frome data (Singh 1981; Singh and Luly 1991). It was argued by Singh and Luly (1991) that there was a change in the balance of summer and winter rainfall reaching Lake Frome with general retreat of summer flowering grasses during dry/cold times. This favours perennial shrublands of Asteraceae and -253-

Chenopodiaceae, adapted to winter rainfall and cold conditions. If these phases at Lake Frome reflect the southward expansion of the monsoon, then wetter conditions in the Flinders Ranges may have occurred (Gell and Bickford 1996).

Increasing proportions of grasses to chenopods were recorded in South-eastern Australia around 12 000-9 000 years B.P and interpreted as a regional increase in rainfall, as this area was outside the seasonal boundary (Kershaw 1995). Further to this, a reversal in the grass:chenopod ratios also occurred, thought to be a response to a reduction in moisture from climatic variability rather than a shift in seasonal rainfall patterns (Kershaw 1995). Previous work on stick-nest rat middens (McCarthy et al. 1996) cautioned that inferring shifts in the summer/winter rainfall boundary on the basis of changes in ratios between grasses and chenopods was simplistic, because the factors affecting plant distributions are complex. In addition to this, the notion of the summer/winter rainfall boundary as a line on a map is simplistic, as it is more a line of transition. Changes in relative abundances of Chenopodiaceae/Amaranthaceae and Asteraceae pollen in the midden records is apparent in Flinders Ranges vegetation communities, with Middle (4 000-2 000 B.P) and Late Holocene (1 000 B.P to present) increases in chenopods and corresponding decrease in Asteraceae at sites in the central ranges. Further, levels of grasses in midden records are relatively stable throughout the Holocene with a slight decrease during the Late Holocene (1 000 B.P to the present) (see Figure 8.3 in Chapter 8). This is contrary to work by Singh and Luly (1991) at Lake Frome, where chenopods and Asteraceae (both winter rainfall taxa) both increased or decreased when grasses (summer rainfall taxa) increased. Behaviour of Asteraceae and Chenopodiaceae/ Amaranthaceae in midden records may be a short-term response to changing availability of total precipitation, rather than associated with longer term shifts in the summer-winter rainfall boundary.

Variability (ENSO) The El Nino Southern Oscillation (ENSO) is recognised as a phenomenon influencing climatic patterns which have impacts on marine and terrestrial environments (Allan 1987). It is suggested that ENSO-induced climatic variability is a feature of the last 5 000-3 000 years while the Early Holocene was less variable (Thorn 1992; McGlone et al. 1992; Schulmeister and Lees 1995; Harberle 1998). The response of vegetation communities in the northern and central ranges from midden records, points to an increase in aridity that occurred after 4 000-2 000 B.P. Higher levels of variability in vegetation communities has been demonstrated midden records after 4 000-2 000 yrs B.P and increasing in the Late Holocene (1 000 B.P to present) middens especially in the northern ranges. Increase in climatic variability in the last 4 000-2 000 yrs B.P as a result of ENSO is supported by evidence from the middens. -254-

Fire Fire as a natural process, may cause a reduction in tree cover and a loss of floristic diversity in vegetation communities (Crowley and Kershaw 1994), the extent of which is influenced by the type and frequency of fire (Kirkpatrick 1994). It can also result in an expansion of fire tolerant vegetation and some communities that have adapted to fire (Hope 1994). Alternatively, fire can maintain plant species diversity in communities (Stafford Smith and Morton 1990) or promote particular plant and animal species in the short term while permanently altering floristic composition over the long term (Hiscock and Kershaw 1992; Hope 1994). However, these impacts are hypotheses requiring further testing, rather than established fact. On a broad scale, climate is the main determinant of vegetation communities, and shifts in the vegetation are largely controlled by precipitation and temperature. Fire is considered a more significant factor at a local scale, as individual taxon can show individual responses to fire (Dodson et al. 1992). For example, potential dominants can be forced out of vegetation communities because of insufficient competitive ability or fire tolerance. Vegetation is sensitive to human activity, especially through fire (Kirkpatrick 1994). However, it is debateable over how much change in vegetation is attributable to human activity versus climate (Hodgkinson 1983; Kirkpatrick 1994; Hope 1994) as it can be difficult to isolate the role of human impact (Macphail 1983; Hodgkinson 1983; Head 1989). Aboriginal burning during the Holocene was used with a high degree of control for purposes of eliminating, creating or shifting of boundaries between vegetation types (Kirkpatrick 1994). It is reasoned that the frequency of fires increased, burning season broadened and intensity declined (Hodgkinson 1983).

Within more temperate areas, fire related impacts are suggested to have increased throughout the Holocene. Since 6 000 years B.P, there has been increased disturbance through fire and human impacts on vegetation (Kershaw 1995). Change in the vegetation, associated with high levels of charcoal in the Lake George (south-eastern Australia) record, saw the expansion of sclerophyll vegetation (decline in Allocasuarina and increase in grasses and Eucalypts) (Dodson et al. 1992). The marked increase in charcoal during the Holocene at Lake George reaches levels not apparent in other records (Hope 1994). Other pollen sequences in this vicinity near Goulburn (Dodson 1986), indicated that Eucalypt woodland and open forest with grassy understorey was dominant since the Early Holocene and that fires were frequent but did not appear to have impacted on a vegetation community that was most likely fire tolerant. There was a considerable degree of local variability evident (Dodson etal. 1992).

Within the .arid and semi-arid zone, the impact of fire has not been tested extensively, although these environments are fire prone. Singh (1981) suggested a relationship -255-

between vegetation changes and fire in the Lake Frome pollen record. However within the changes, was a climatically influenced component. Fire was considered as an important role in developing the essential character of arid land vegetation during the last 10 000 years (Luly 1993). During the Early Holocene (8 000-7 000 B.P) there was evidence of an increase in fire from charcoal levels. Woodland, shrubland and grasses declined, while ephemeral taxa such as Asteraceae increased. During 7 000-4 200 B.P there were short fire phases and trees began to recover. An increase in charcoal from 4 200-2 200 B.P saw a rise in ephemerals and decline in trees. From 2 200 B.P to the present, fires were present (especially after 650 B.P), there was an increase in chenopods from the dominance of winter rainfall, and a slight increase in trees and shrubs (Singh 1981; Singh and Luly 1991).

Records from Lake Tyrrell suggested that fire effects were minimised in the vegetation during the Middle Holocene (6 600 B.P) and grasses were out competed by Callitris woodlands (Luly 1990; Luly 1993). Drier conditions intensified after 2 500 B.P when fire caused the attrition of Callitris and expansion of grasses in the more open vegetation community (Luly 1990). A drastic decrease in tree cover and rise in grasses and charcoal indicated the arrival and impact of European agriculture (Luly 1993).

The types of changes in vegetation caused by fire from these examples in the literature is not evident in the vegetation records from Leporillus middens. The resolution of reconstructed communities from middens is broad scale and middens provide no independent evidence of fire (ie. charcoal), unless they are burnt. The retreat of woodland and shrubland communities and increase in chenopod shrublands, is a likely response to climatic change. Within the semi-arid context, regular burning of woodland communities maintain a dense grass layer (Hodgkinson 1983). Prevention of grass fires in semi-arid woodlands causes an increase in the density of shrubs and replacement of long lived perennial grasses with short lived grasses. Fire kills much of the shrub layer in chenopod shrublands and regeneration by seed is slow, while herbs regenerate quickly (Hodgkinson 1983). Changes in floristic composition and/or boundaries of fire sensitive and fire tolerant communities caused by fire, are not detected in the midden records. It should be acknowledged that the increase in variability in vegetation records from the Late Holocene middens (1 000 B.P to present) is consistent with increased fire and instability, even though on the basis of the midden evidence, these factors cannot be tied in directly.

Topography, moisture availability and soil nutrient levels Factors important in the functioning of arid environments have been presented in a conceptual model by Stafford Smith and Morton (1990). The factors that affect plant communities in the model are discussed within the context of the Holocene vegetation -256-

records from middens, to explore the extent to which they may be responsible for the changes that have been observed. The spatial heterogeneity of arid environments produces patterns in the floristic composition and structure of vegetation communities (Stafford Smith and Morton 1990). This heterogeneity is caused by unpredictable and extreme rainfall events, soil nutrient and moisture levels and the diversity of life histories adopted by plant taxa (eg. ephemeral and perennial species).

Rainfall has a high temporal variability (annual and seasonal rainfall) across different regions in arid and semi-arid environments. Large rainfall events can recharge water tables, increase soil moisture and trigger the growth of long lived perennial taxa. This is compounded by the fact that variable topography results in different amounts of water and nutrients in runoff, that in turn affects patterning of plant assemblages that are often separated by abrupt boundaries (Stafford Smith and Morton 1990). Extremes of soil moisture are crucial in defining diversity of plant life histories on different substrates. Rugged topographic settings receive relatively higher levels of water and nutrients in runoff and thus tend to be dominated by long lived perennial plants (eg shrubs). These taxa can establish large root systems, making them more tolerant of drought. Areas with intermittent water supplies (eg floodplains) carry the widest range of taxa that have adopted different life history strategies (ephemerals, short lived perennials and long lived perennials that are opportunistic). Areas with a continuous water supply are dominated by perennial taxa in the absence of disturbance such as grazing or fire. Less diverse topographic settings have low levels of soil moisture and nutrients. These conditions encourage faster growing plants that use nutrients quickly but are less drought tolerant (Stafford Smith and Morton 1990).

The northern and central Flinders Ranges is a region of diverse topography and habitats that have produced a mosaic of semi-arid vegetation communities on the landscape. The northern ranges are more rugged, with a relatively higher rainfall compared to the central ranges. Therefore, availability of moisture from runoff and nutrient levels are higher, providing conditions for the establishment of long lived tree and shrub taxa. The northern ranges support a higher density of plant species than the central ranges (Greenwood et al. 1989) and a greater occurrence of relict flora has been recorded. For example, Melaleuca uncinata, Calytrix tetragona and Eriostemon augustofolia usually found in southern Australia in heath communities on sandy substrate, are found in the Arkaroola-Mount Painter Sanctuary. There are also trees including Capparis mitchelli, Acacia confluens and Codonocarpus pyramidalis (Sprigg 1984). The central ranges have a less extensive rugged topography, confined to gorges that transverse rolling hills and plains, and receive less rainfall. Open shrublands, chenopod shrublands and herb fields are common communities in these environments. Relict flora have been recorded -257-

for example, Capparis mitchelli, Santalum acuminatum and Melaleuca dissitiflora, confined to main creeks in Mount Chambers Gorge (Medlin 1993), but are less common than in the northern ranges.

The changes in dominance of woodland/shrublands and chenopod shrublands during the Holocene, inferred from Leporillus spp. middens are to some degree, an artefact of characteristics of semi-arid zone ecology, as environmental conditions in the central ranges are predisposed to dominance of chenopod shrublands while the north favours topographically buffered woodland and shrubland communities that more commonly include relict species. Therefore, these biogeographical parameters are responsible for maintaining the variability that is evident in vegetation records from the youngest middens. The northern ranges may well be less sensitive to climatic change, compared to the central ranges, given the environmental characteristics as previously outlined. These biogeographical parameters do in part explain the less dramatic shift in vegetation communities throughout the Holocene that was observed in midden records from the Arkaroola-Mount Painter Sanctuary in the north, as opposed to the central ranges.

9.3 Holocene Palaeoclimates in the Flinders Ranges

This current study agrees with the interpretation of wetter conditions with dominance of summer rainfall until 6 000 years as evidenced by pollen and macrofossil records from the middens at Brachina Gorge and Mount Chambers Gorge, as woodlands were more widespread than present. Suggestion of a decrease in total rainfall from 4 500-2 200 years B.P (with winter rainfall a certain occurrence), is supported by middens from the central ranges as chenopod shrublands became more widespread. However in the northern ranges at this time, woodland and shrublands persisted. Rugged terrain in the northern ranges and more effective precipitation available are probable causes for maintaining woodland and shrubland communities in a climate that was becoming more arid. It is difficult to reconcile Late Holocene midden records with Singh and Luly's (1991) suggestion that from 2 200 years to the present there has been a slight increase in summer rainfall. Chenopod shrublands have increased over this period in the midden records from the central ranges and remained largely unchanged in the north, while changes in grass xhenopod ratios were not apparent in the midden records. The striking feature of Late Holocene midden records was the increased spatial variability in vegetation records that began after 4 000-2 000 yrs B.P, especially in the northern ranges and was most prominent from 1 000 yrs B.P to the present. Interestingly, topographically buffered vegetation in the north, shown to be more resilient than vegetation from the central ranges to climate change, has implications for ENSO driven climatic variability and the relationship between short and long term change. -258-

9.4 Conclusion

Timing and direction of change in Holocene vegetation in the Flinders Ranges corresponded closely to controls exerted by climate. Early to Middle Holocene (7 000-5 000 B.P) conditions provided a suitable environment for woodland and shrubland communities on the landscape in both the northern and central ranges. As the climate was becoming increasingly more arid during 4 000-2 000 B.P, causing a reduction in total precipitation and thus less soil moisture and nutrient levels, woodlands and shrublands with herbaceous understories from the central ranges were not suited to the changing conditions. The vegetation adapted by shifts in the abundance of whole communities, resulting in more widespread and dominant chenopod shrublands. Changes in the northern ranges were not as dramatic, given the existing landscape characteristics, and floristically diverse woodland and shrubland communities were maintained during this part of the Holocene. Late Holocene (1 000 B.P to present) vegetation communities display high levels of variability, that support arguments for increased variability of significant climatic parameters such as ENSO, during the Late Holocene. -259-

Chapter 10: Conclusions and Further Work

The research undertaken in this thesis was designed to understand the taphonomy of Leporillus middens and provide a systematic regional study of Holocene vegetation histories and palaeoclimates in the central and northern Flinders Ranges, South Australia. The modern spatial variability of the Flinders Ranges vegetation, and diverse habitats in a rocky upland environment within the semi-arid zone, presented important implications for interpretation of midden pollen and macrofossil records.

Modern pollen rain at study sites was investigated to provide the foundation for interpretation of pollen from the stick-nest rat middens. Regional pollen rain from west- east transect studies in the central and northern Flinders Ranges reflected the high spatial variability in vegetation communities from semi-arid rocky upland environments. Different vegetation communities throughout the ranges (eg. riverine woodlands, tall shrublands and chenopod shrublands) could be identified in the modern pollen spectra. Temporal variability in the modern pollen rain was detected along the west-east transects in both the northern and central ranges. Local environmental conditions at midden cave sites, including cave aspect and degree of exposure, and composition of local vegetation cover, were significant factors in determining the composition of pollen rain at midden sites. Pollen traps located inside and outside caves recorded strong signals from local taxon and less dominant signals from regional pollen types.

Variability in midden palaeoecological records may be influenced by processes other than vegetation change over time. Of particular concern in this research, was changing patterns of pollen representation with changes in source areas at different midden sites, and the influences of rat activity. To separate these processes, chronology of middens was investigated to provide insight into the taphonomy of deposits and the possible influences of the rats. Fossil pollen records from middens were also compared with modern vegetation and modern pollen rain and fossil pollen and macrofossils were compared.

Midden deposits accumulate fairly rapidly, as evidenced by multiple AMS dates on sub samples of single midden deposits, and therefore a representative sample from an extensive midden is likely to return a reliable age for the deposit. In most instances, ages of faecal pellets and leaf macrofossils were contemporary when excluding possible systematic discrepancies from different dating laboratories. However, it is acknowledged that reworking of material may result in different ages of faecal pellets and leaf macrofossils from the same sample. Dating pollen concentrates was more problematic. In most cases, the age of pollen corresponded with ages of macrofossils from the same sample. Pollen dates should be regarded as imprecise but reliable given that midden -260-

pollen used to reconstruct vegetation, represented an averaged record for the relevant sub- sample of midden.

Pollen recruited into midden assemblages is a mix of local taxa from outside cave sites and regional taxa from outside the immediate environment. MacrofossihPollen Index (MPI) values indicated that macrofossils are a secondary source of information in Leporillus spp. middens, given the low relative abundance and diversity of macrofossil taxa and variable quality of preservation. MPI values have shown that pollen taxa in middens are not over represented as a result of Leporillus spp. activity, rather that changes in the pollen abundance reflect real vegetation change.

There were spatial and temporal variations in midden vegetation records from well dated sites from two study areas in the central ranges and one study area in the northern ranges. Early Holocene (7 000-5 000 B.P) and Middle Holocene (4 000-2 000 B.P) vegetation records from sites in the central ranges were similar, while after 4 000-2 000 yrs B.P and into the Late Holocene (1 000 B.P to present), records reflected the spatial variability of vegetation communities from present day semi-arid environments. Similar trends were observed in records from midden sites in the northern ranges.

Change in Holocene vegetation communities was more apparent in the central ranges as there was a shift from open woodland and shrubland communities with herbaceous understories at 7 000-5 000 B.P, to chenopod shrublands in the Middle (4 000-2 000 B.P) and Late Holocene (1 000 B.P to present). Woodland and shrubland communities with herbaceous and grassy understories were more extensive in the northern ranges during the Early Holocene (7 000- 5 000 B.P). Less extensive coverage of both communities persisted into the Middle (4 000-2 000 B.P) and Late Holocene (1 000 B.P to present), with a slight increase in chenopods and decrease in grasses in the understorey.

Vegetation records across the Flinders Ranges at 7 000-5 000 B.P are a response to increased effective precipitation and accord with other Holocene palaeoclimatic records suggesting wetter and warmer conditions than the present. Retreat of woodland and shrubland communities and consequent expansion of chenopod shrublands in the central ranges are indicative of drier and cooler conditions with increasing aridity, that has also been recorded in other evidence from around the continent. Shifts in vegetation communities as a climatically driven response, was more visible in the central ranges, as a result of different environmental conditions more sensitive to changes in effective precipitation. This highlights the complexity of factors that affect plant distributions in semi-arid environments. Underlying the climatic driven response is the role of biogeographic parameters that influence the diversity and structure of different vegetation -261-

communities. In sheltered rugged topography, characteristic of the northern ranges, plant communities are more stable as evidenced by the occurrence of relict species and maintenance of woodland and shrubland communities throughout the Holocene, even as conditions were becoming increasingly arid. The same climatic scenario of increasing aridity in the central ranges resulted in less stable vegetation communities that responded to cooler and drier conditions by shifting from dominantly shrublands to extensive chenopod shrublands and herblands after 4 000- 2 000 yrs B.P. Present spatial variability in the vegetation being a feature of the last 1 000 yrs (and possibly longer in the central ranges) with more homogeneous conditions across the ranges from 7 000-5 000 yrs B.P, is certainly consistent with the argument that climatic parameters such as ENSO have become more variable during the last 4 000-3 000 years.

The argument that topographically buffered vegetation from the northern ranges is more resilient than other communities from the central ranges to climate change, raises some interesting questions for future research. Findings from this research has important implications for understanding the palaeoecology of refugia in relation to debates about human occupation, flora, and fauna within the arid zone. Quantifying the extent of change in vegetation also has important implications for increasing our understanding of arid zone ecology and the relevance of a long-term ecological perspective for future management of our fragile semi-arid and arid lands. Increasing our understanding of the complex interactions between short and long-term climatic and biogeographic change in the semi-arid zone is crucial for a significant part of the continent that forms a broad transition zone between the arid inland and coastal fringe of Australia. -262-

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Wilson, A. (1990) The effect of grazing on Australian ecosystems (In) D. Saunders, A. Hopkins and R. How (Eds.), Australian Ecosystems 200 years of Utilization, Degradation and Reconstruction, Surrey Beatty and Sons, Sydney, Pg 235-244.

Winkworth, R. E. and Thomas, P. R. (1974) Climate, Department of Primary Industries, Queensland.

Wyrwoll, K. H., Hopwood, J. and McKenzie, N. L. (1992) The Holocene Paleohydrology and Climatic History of the Northern Great Sandy Desert-Fitzroy Trough: with Special Reference to the History of the Northwest Australian Monsoon, Climatic Change 22: 47-65.

Wyrwoll, K.H., McKenzie, N.L., Pederson, B. and Tapley, LB. (1986) The Great Sandy Desert of Northwestern Australia: the Last 7 000 years, Search 17:208-210. -283- APPENDIX 1: Weights of middens and >2mm and >500um sorted fractions

Midden Weight (prior soaking) (g)Weigh t (>2mm) (g) Weight (>500um) (g) Total (g)

BR1 TOP 1089.1 211.75 211.75 BR1 MIDDLE 433.3 63.7 2.05 65.75 BR1 BASE 394.7 41.44 41.44

BR3TOP 404.99 139.5 65.23 204.73 BR3 MIDDLE 318.6 53.5 1.06 54.56 BR3 BASE 174.7 19.25 8.25 27.5

BR4 (i) 272.4 74.6 74.6 BR4 (iii) 198.9 47.5 47.5 BR4 (iv) 81.9 79.2 79.2 BR4 (v) 61.3 59.5 59.5 BR4 (vi) 128.96 43 43 BR4 (vii) 232.3 56.8 56.8

BR7 TOP sample 1 586.3 62.8 62.8 BR7 TOP sample 2 467.7 56 56 BR7 MIDDLE 679.61 149.05 6.93 155.98 BR7 BASE 371.6 34.9 9.7 44.6

BR2 718.3 50.1 22.4 72.5

HH1 TOP 528.4 37.03 37.03 HH1 BASE 316.4 33.22 33.22

NWCK1 474.2 123.2 87.75 210.95

NWCK2 590.41 117.09 117.09

WF1 342.7 90.23 90.23

RC1 sample 1 369.94 99.5 99.5 RC1 sample 2 555.6 121.56 121.56

RC2 387.3 154.78 182.07 336.85

RC3TOP 223.19 36.9 0.42 37.32 RC3BASE 109.08 26.93 26.93

ARK1 TOP 89.05 29.59 29.59 ARK1 MIDDLE 52.28 4.76 4.76 ARK1 BASE 33.04 30.04 30.04

OT1 sample 1 227.5 221.61 221.61 OT1 sample 2 274.13 74.5 74.5

MC1 EXPOSED TOP 172.9 17.85 17.85 MC1 EXPOSED BASE 184.9 23.96 23.96

MC1 TOP 255.7 64.48 64.48 MC1 INNER CENTRE 11.94 11.94 MC1 BASE 444.3 56.75 56.75

MC2TOP 338.7 32.31 32.31 MC2TOP&OVERHANG 243.5 76.84 76.84

CG sample 1 268.4 102.27 102.27 CG sample 2 i 672.4 39.69 39.69

MD3 524.4 64.06 9.28 73.34 -284-

APPENDIX 2 : Standard Method for Pollen Extraction

Initially, fifty sub-samples of midden matrix suspended in amberat solution underwent the traditional alkali treatment. Approximately 5cm^ of samples were placed in a 10% solution of Potassium Hydroxide (KOH). This breaks up the structures of plant remains and removes lignites and humic acids. Boiling the samples in KOH for 20 minutes produced a dark brown supernatant composed of dissolved organic acids. This solution was then sieved through a 100 |J,m mesh into centrifuge tubes to remove sediment and large organic particles. Samples were then placed into Hydrofluoric Acid (HF) and left for forty eight hours. This ensured that all silica was dissolved. They were then balanced with a 10% solution of Hydrochloric Acid (HC1) to remove any silico- fluorides in solution, centrifuged, decanted and washed with distilled water.

The pre-acetolysis procedure followed this step. Samples were transferred to glass testubes and dehydrated by successive washes in 30% acetic acid and glacial acetic acid. Dehydrated samples were acetolysised, to hydrolyse cellulose and other polysaccharides into water soluble products such as glucose. This consisted of placing samples into Acetic Anhydride and 10 drops of Sulphuric Acid (H2SO4), heated for 1 minute while stirring the mixture continuously. This strong oxidation reaction removed organic material and darken pollen grains to make features visible when viewed under the light microscope.

To prepare pollen concentrates for mounting on a microscope slide, samples were rehydrated by washes in glacial acetic acid, 30% acetic acid and distilled water. Samples then underwent an alcohol series by washing in 80% ethanol, centrifuge and decant, followed by 100% ethanol, centrifuge and decant and finally Tertiary Butyl Alcohol (TBA) transferred to glass phials, decanted and capped. Silicon oil was added to the pollen concentrates to ensure pollen concentrates were stored without being dehydrated or altered. Silicon oil is recommended as it has a low refractive index where pollen grains are clearly visible and mobileand can be examined from different angles in the identification process (Moore and Webb 1983). -285-

APPENDIX 3 : Features of Pollen Grains For Identifications

Acacia Description: grains in polyads with a psilate or granulate exine. Size: 40 - 100 um Major likely sources: Acacia victoriae, A. rivalis, A. tetragonophylla, A. salicinia, A.confluens, A. ligulata.

Apiaceae Description: tricolporate grains thickening exine in the sub polar region. Oval shape with long colpi and medium to large sized grains. Size: 20 - 30 um Major likely sources: Trachymene glaucifolia, Daucus glochidiatus, Hydrocotyle spp.

Asteraceae Description: tricolporate oblate-spheroidal to prolate echinate with well developed spines on the surface. Size: 15 -100 um Major likely sources: Chryscephalum spp., Brachycome spp., Cassinia spp., Olearia spp., Ixolaena spp., Pluchea spp.,Vittadinia spp.

Boraginaceae Description: Mostly rectangular shape and rounded at the edges. Colporate or colpi cross shaped and the exine is thin and psilate. Size: 12 - 20 um Major likely sources: Echium plantagineum, Trichodesma zeylanicum

Caesalpinaceae Description: Trizonocolporate with small often indistinct pores that tend to be apertures at the equatorial constriction of the long colpi. Grains are subprolate to spheroidal and surface can be granular and are susceptible to crushing. Size: 30 - 50 um Major likely sources: Senna artemisioides ssp. -286-

Casuarinaceae Description: Oblate to apiculate in lateral view and rounded. Triporate and pores are small and angular and the exine at the apertures is thickened. Surface of the grain is psilate to scabrate. Size: 20-30 um Major likely sources: Casuarina cristata, C. cristata spp. pauper, C. verticulata

Chenopodiaceae/Amaranthaceae Description: Monad spherical grain and polyporate with small pores. Three groups identified on the basis of the number of pores from a lateral view. Chenopod 1: 3 pores, Chenopod 2: 6 pores and Chenopod 3: 9 pores Size: 12 - 40 um Major likely sources: Ptilotus spp., Atriplex spp., Maireana spp., Rhagodia spp., Chenopodium spp., Enchylaena spp., Sclerolaena spp. and Salsola spp.

Convolvulaceae Description: Large spheroidal to subprolate grains with a thick exine that is psilate. Colpi are sunken. Size: 50 - 80 um Major likely sources: Convolvulus remotus, Convolvulus erubescens.

Cruciferae Description: Tricolpate and variable grain size. Grains are spheroidal to subprolate with a coarse reticulate exine. A distinctive lobed profile view. Size: 15-30um Major likely sources: Sisymbrium spp., Carrichtera spp.

Cupressaceae Description: Spherical grain, inaperturate with a thin exine. Size: 20 - 30 um Major likely sources: Callitris columellaris

Cyperaceae Description: Elongated grain with one end broader than the other. One pore at the broad end and three at the other. Very thin exine. Size: 20 ->60 um Major likely sources: Cyperus spp., Schoenoplectus spp. -287-

Epacridaceae Description: Tetrads with monad spherical gains. Each grain are tricolporate with transverse furrows Size: Tetrad diameter 40 - 50 um Major likely sources: Epacris spp.

Eucalyptus Description: Syncolpate thickened exine near the apertures and a polar island Size: 10 - 15 um Major likely sources: Eucalyptus camaldulensis, E. intertexta, E. obtusa, E. socialis, E. gillii, E. microtheca.

Euphorbiaceae Description: Tricolporate, prolate, subprolate to oblate spheroidal grains. Long colpi and pori not always seen. Exine is psilate with distinct columellae. May form a reticuloid pattern. Major likely sources: Euphorbia spp.

Fabaceae Description: Trizonocolporate subprolate grain with a reticulate exine. Triangular in polar view with narrow long sunken coplistriate-rugulate surface. Size: 20 - 30 um Major likely sources: Glycine spp., Indigofera spp., Clianthus formosus

Geraniaceae Description: Tri-colporate polyforate or inaperturate short colpi and rounded pori. Grains can be large, prolate-spheroidal to suboblate with sculptering on the exine in the form of striations. Size: 40 -100 um Major likely sources: Erodium spp.

Goodeniaceae Description: Tricolporate prolate oval/spheroidal grain. Granulate or tectate elevate and smooth around the furrows. Distinctive tectate exine. Size: 25 - 30 um Major likely sources: Goodenia vernicosa, Scaevola spinescens. -288-

Haloragaceae Description: Grains are paraisopolar and radially symmetrical with 4-5 porate. The exine can be psilate and thin to thick and granulate. Size: 12 - 30 um Major likely sources: Haloragis spp., Gonocarpus spp.

Lamiaceae Description: Tri-colpate oblate to prolate grain and usually reticulate. Major likely sources: Prostanthera striatiflora, Marrubium spp.

Leptospermum Description: Syncolpate oblate with concave sides. A triangular outline from the polar view. Size: 12-18 um Major likely sources: Leptospermum laevigatum, L. coriaceum, Ljuniperinum.

Liliaceae Description: Ellipsoid grain and monosulcate. Sise: 30 - 70 um Major likely sources: Arthropodium spp., Dianella spp., Bulbine spp. Xanthorrhoea spp.

Loranthaceae Description: Tricolpate with long narrow colpi. Grains are peroblate to subprolate and triangular. Sides are convex or concave and the surface appears stippled and smooth. Size: 30 - 70 um Major likely sources: Lysiana spp.

Lythraceae Description: Isopolar radiosymmetrical grain with 3 long colpi from a pore. Subprolate with a thin exine Size: 30 - 40 um Major likely sources: Lythrum spp.

Malvaceae Description: Polyporate suboblate/oblate spheroidal and moderately large with short spines and a granulate background. 30 to >100 um Major likely sources: Abutilon spp., Sida spp. -289-

Melaleuca Description: Small oblate grains with straight sides in polar view. Syncolpate (or parasyncolpate). Size: 12.5-22.5 um Major likely sources: Melaleuca glomerata

Myoporaceae Description: Tricolpate prolate or oblate-spheroidal grain. Elliptical colpi with one aperture above and below the equator Size: 20 - 25 um Major likely sources: Eremophila spp., Myoporum spp.

Onagraceae Description: Triporate grain usually large peroblate to spheroidal with a granular exine. Triangular in polar view with protruding pores. Size: 45 - > 120 um

Other Myrtaceae Description: Small to medium size grains tri-colporate with triangular polar view and smooth exine. Parasyncolpate grains that may or may not have an island. Size: < 40 um Major likely sources: Baeckia spp., Calytfix spp., Kunzea spp.

Pinus Description: Spheroidal grains with 2 air bladders Size: Major likely sources: Pinus radiata

Pittosporaceae Description: Tricolporate prolate - spheroidal and small to medium sized grains. Size: 20 um Major likely sources: Bursaria spinosa

Plantaginaceae Description: Spheroidal and periporate. Major likely sources: Plantago spp. -290-

Poaceae Description: Spherical to elliptical grains with single protruding pore and distinct annulus. Thin exine that is usually smooth/psilate. Size: 20 - 30 um Major likely sources: Cymbopogon ambiguus, Stipa spp., Danthonia spp., Enneapogon spp., Aristida spp., Eriachne, spp., Digitaria spp., Paspalidium spp., Themeda spp., Triodia spp.

Polygalaceae Description: Polycolporate and grain is suboblate to prolate. Size: 30 um Major likely sources: Comesperma spp.

Polygonaceae Description: Smooth tricolporate grain with flat narrow colpi. Major likely sources: Acetosa vesicarius

Portulacaceae Description: Variable morphology, usually tricolpate and panotreme with apertures being transistional between colpi and pores. Major likely sources: Calandrinia spp., Portulaca spp.

Proteaceae Description: Triporate grain with concave sides narrowing towards the pores. Oblate to oblate-spheroidal and large. Thick scabrate exine with distinct costa and annula. Size: 50 - 80 um in the polar view Major likely sources: Hakea leucoptera, H. edneiana, Grevillia spp.

Rutaceae Description: Variable shape and size tricolporate grains with a psilate exine. Grains are suboblate to subprolate. Size: 15 -100 um Probable Sources: Eriostemon spp., Boronia spp. -291-

Santalaceae Description: Tri-colporate subprolate small grains with a long and narrow colpus. Other grains can be triporate prolate, with rounded apertures. Can appear triangular in polar view. Size: 10 - 45 um Major likely sources: Exocarpus spp., Santalum spp.

Sapindaceae Description: Circular-semilobate/prolate oval grain. Psilate exine with large pores and distinctive costa. Some grains have bulbous protrusions between the furrows in the polar view. Size: 18-30um Major likely sources: Dodonaea spp., Alectryon spp.

Solanaceae Description: Tri-colporate circular/spherical grain with thin psilate exine and protruding costa. Size: 15 - 20 um Major likely sources: Solanum spp.

Stackhousiaceae Description: Tricolporate grains that are reticulate and oblate/spheroidal or prolate- spheroidal. Circular comparatively large pores. Size: 10 - 40 um Major likely sources: Stackhousia spp.

Thymelaceae Description: Polyporate spheroidal grain with an exine displaying a croton pattern. Size: 20 - 25 um Major likely sources: Pimelea spp.

Typhaceae Description: Single monoporate grains but can occur as tetrads. Round pore with no annulus and an exine which is finely reticulate. Size: 30 - 35 um Major likely sources: Typha domingensis -292-

Zygophyllaceae Description: Tricolporate grain with circular pores, subprolate and finely reticulate. Size: 15 - 35 um Major likely sources: Zygophyllum spp., Nitraria spp.

Appendix 3: Summary of pollen characteristics used for identification of pollen in middens and pollen traps. Descriptions have been compiled from Boyd (1992) and likely major sources from species inventories collected in field work and flora lists from Greenwood et al. (1989), Cunningham et al. (1992) and Gell and Bickford (1996). APPENDIX 4: Pollen Counts for Middens and Traps

Midden pollen counts Pg 293-300 1995 and 1996 pollen trap counts from west-east transect study Pg 301-306 1995 and 1996 pollen trap counts from midden cave sites pg 307-318 -293-

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CO 10 co CO CD CD CD CD 5 CD CD 2 2 1— 4— cP r-

co LO CO LO CD CD CO 3 CO CO CO 4— o o CO CO 8 CO o o> o a 3 3 g 3 o 8 T— 3 8 o. 8 T— 3 8 _3 3 o co O 3 O CO CO CD co o s o CM CO CM o I CO o to LL to Q. CM T— •o o. a a a CM CD o. LL o T- Q o TJ Q CO CO o 8 to CO CO o O O co O to h- o k_ O CD Q 1- LL o ca h- E 2 o 2 2 2 5 CO 2 2 2 H 2 1 Ul o 5 o 2 ca Ul o o -327-

Tt CM CO o CD CM CO CM co o p co o co o o

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APPENDIX 6 : Preparation of Pollen Concentrates for Accelerator Mass Spectrometry (AMS) Radiocarbon Dating

Pretreatment of 5 cm^ sediment samples for pollen concentration after Brown et al. (1989):

1) Samples boiled in 6% KOH for 20 minutes and sieved at 180|xm 2) The <180|im fractions treated with 48% HF in boiling water bath for 25 minutes and then with 1 N HC1 in boiling water bath for 10 minutes. 3) Pretreated residues sieved at 88|im and <88(im fraction bleached with 2-3% NaOCl for 5 minutes 4) Fraction sieved at 44|nm* and retain the 44-88(xm fraction. 5) The <44uLm fraction rebleached with NaOCl for 5 minutes and sieved at 20(im retaining the 20-44(im fraction. Discard the 20u\m fraction. 6) The retained fractions (44-88|im) and (20-44|a.m) pollen concentrates are AMS dated

* Sieving mesh sizes can be changed and are selected to eliminate organic matter both larger and smaller than most pollen grains (Cwynar et al. 1979).

Pretreatment of 15 cm^ sediment samples for pollen concentration after Regnell (1992):

1) Place sample in hot 10% NaOH for 15 minutes and hot 10% HC1 for 10 mins. 2) Sieve the sample at 40|im 3) < 40|im fraction retained and placed in hot 48% HF for 20 minutes 4) Repeat the hot 10% HC1 treatment for 10 minutes 5) Sieve the sample at 20|j,m 6) < 20(im fraction retained and bleached in 2% NaOCl for 5 minutes 7) Place in concentrated H2SO4 for 3 hours 8) Sieve at 10fj.m 9) AMS date the 10-20|4,m fraction.

The Sodium Hypochloride (NaOCl) treatment step by Brown (1989) was altered by adding Sulphuric Acid (H2SO4) to the procedure to remove organic detritus. Polysaccharides are hydroloysed by the H2SO4 and the treatment does not appear to affect the exines of pollen grains. NaOCl destroys lignin but is also corrosive to pollen grains unless in a weak solution and exposed to grains for a short period of time (Regnell 1992). -329-

APPENDIX 7: Comparison of 1995 and 1996 West-East Transect Pollen Trap Studies at Arkaroola, Mount Chambers Gorge and Brachina Gorge

I

Clustering History Number of Clusters Distance Leader Joiner 6 5.5560254324 A/1:1995 A/23:1995 5 5.8296879864 A/10:1995 A/1:1996 4 6.9278299692 A/10:1995 A/10:1996 3 7.4508750404 A/10:1995 A/11:1995 2 8.9678893041 A/1:1995 A/14:1996 1 9.9605177504 A/1:1995 A/10:1995 Dendrogram

A/1:1995 A/23:1995 A/14:1996 A/10:1995 A/1:1996 A/10:1996 A/11:1995

Appendix 7(1): Comparison of all pollen taxa from sites along the Arkaroola-Mount Painter Sanctuary west-east transect for the 1995 and 1996 sampling period (less other myrtaceae, exotics and unknowns). -330-

II

Clustering History Number of Clusters Distance Leader Joiner 7 5.069036078 C/12:1996 C/13:1996 6 6.3423479176 C/11:1996 C/12:1996 5 6.5475302549 C/9:1996 C/10:1996 4 7.7312474084 C/9:1996 C/11:1996 3 8.3419879969 C/9:1995 C/9:1996 2 8.5919925907 C/9:1995 C/13:1995 1 10.393558427 C/9:1995 C/11:1995 Dendrogram

C/9:1995 0 C/9.1996 C/10:1996 I 1 C/11:1996 C/12:1996 C/13:1996 !- C/13:1995 C/11:1995 0

Appendix 7(11): Comparison of all pollen taxa from sites along the Mount Chambers Gorge west-east transect for the 1995 and 1996 sampling period (less other myrtaceae, exotics and unknowns). -331-

III

Clustering History Number of Clusters Distance Leader Joiner 5 7.0364257584 B/8:1996 B/3:1996 4 7.1254756844 B/9:1995 B/8:1995 3 7.7091772676 B/2:1996 B/4:1996 2 7.9426165296 B/8:1996 B/2:1996 1 9.8589927142 B/9:1995 B/8:1996 Dendrogram

B/9:1995 B/8:1995 B/8:1996 B/3:1996 B/2:1996 B/4:1996

Appendix 7 (III): Comparison of all pollen taxa from sites along the Brachina Gorge west-east transect for the 1995 and 1996 sampling period (less other myrtaceae, exotics and unknowns). APPENDIX 8: Chi-Square Statistics for Pollen Trap Data -332-

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Mt . VO vn 00 Tt Ov VO 00 in oo en en CN OO Chamber s Wes t Os CM CN en VO o r- T-H CN Tt

so Os CN O m O 00

Gorg e en Brachin a in en OS CN Tt cn i-H

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HH l CN NWC K Os CN O en CN CN CM & m SO 00 Tt Os Tt 4— O Os CN 1 1 en in HH l NWC K O in VO Os CN CN

VO VO Os m in 00 V, 4— VO HH l 00 CN Wes t Os Tt 4-H m © in Ov en VO in Os OS 00 © m 00 en CN

HH l 4— Wes t Os c Tt CN en CN r-4 CA CA eu o •o *4-4 CO CA ca CA o "8 CA (U eu CA C/3 X CA 8 ca eu % 3 kH O 4H 4-4 IH CA on o c > CN 1 i eu X X O -335-

APPENDIX 9: Macrofossil:Pollen Index (MPI) Values for Key Taxa in Leporillus spp. Middens

Relative abundance scale for macrofossils (after O'Rourke and Mead 1985) and pollen (afte Anderson and VanDevender 1991): macrofossils relative abundance pollen <5% 1 <1% 5-10% 2 1-9% 11-50% 3 10-24% 51-79% 4 25-49 >80% 5 >50%

MPI=S(Mi-Pj)/N where M= relative abundances of macrofossils and P= relative abundances o pollen for each taxon (i) in middens. N= number of samples. Chenopodiaceae/ Am.aranthaceae Asteraceae Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) M-P WF1 0 3 -3 0 3 -3 NWCK1 1 2 -1 1 2 -1 NWCK2 0 3 -3 0 4 -4 RC1 1 2 -1 1 4 -3 RC2 1 3 -2 0 3 -3 RC3 TOP 0 2 -2 1 3 -2 RC3 BASE 1 2 -1 1 4 -3 ARK1TOP 0 2 -2 1 4 -3 ARK1MID 0 3 -3 1 3 -2 ARK1 BASE 1 3 -2 2 5 -3 HHl TOP 0 3 -3 0 3 -3 HHl BASE 0 3 -3 0 3 -3 OT1 1 5 -4 0 2 -2 N= 13 -30/13 -35/13 MPI -2.3 -2.7

Malvaceae Acacia Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) M-P WF1 0 1 -1 0 1 -1 NWCK1 0 1 -1 1 1 0 NWCK2 1 1 0 1 1 0 RC1 1 2 -1 0 1 -1 -336-

RC2 0 1 -1 1 2 RC3 TOP 2 -1 1 1 RC3 BASE 2 -1 1 1 ARK1TOP 1 0 0 1 ARK1MID 2 -1 0 1 ARK1 BASE 1 0 0 2 HHl TOP 0 1 -1 0 1 HHl BASE 0 2 -2 0 1 0T1 0 1 -1 1 1 N= 13 -11/13 -9/13 MPI -0.85 -0.85 -0.70

Myoporaceae Poaceae Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) WF1 0 2 -2 0 2 NWCK1 0 2 -2 1 2 NWCK2 0 1 -1 0 2 RC1 0 3 -3 1 2 RC2 0 1 -1 1 2 RC3 TOP 1 2 -1 1 2 RC3 BASE 1 2 -1 0 2 ARK1TOP 0 2 -2 0 2 ARK1MID 0 2 -2 0 2 ARK1BASE 1 2 -1 0 2 HHl TOP 0 2 -2 0 2 HHlBASE 0 2 -2 0 2 OT1 1 2 -1 0 1 N=13 -21/13 -21/13 MPI -1.61 -1.61

Casuarinaceae Apiaceae Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) WF1 0 2-2 0 2 NWCK1 0 1 -1 1 1 NWCK2 0 2-2 0 2 RC1 1 1 0 1 2 RC2 0 2-2 1 2 RC3 TOP 0 1 -1 1 3 -337-

RC3 BASE 0 1 -1 0 3 ARK1 TOP 0 1 -1 1 1 ARK1MID 0 1 -1 0 2 ARK1 BASE 0 1 -1 0 2 HHl TOP 0 2 -2 0 3 HHlBASE 0 1 -1 0 3 0T1 1 1 0 0 2 N= 13 -15/13 -23/13 MPI -1.15 -1.77

Sapindaceae Caesalpinaceae Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) WF1 0 2 -2 0 1 NWCK1 0 1 -1 1 1 NWCK2 0 2 -2 0 2 RC1 0 2 -2 0 2 RC2 0 2 -2 0 2 RC3 TOP 0 3 -3 0 2 RC3 BASE 1 3 -2 1 2 ARK1TOP 1 1 0 0 2 ARK1MID 0 1 -1 0 2 ARK1 BASE 1 1 0 1 2 HHl TOP 0 2 -2 0 2 HHl BASE 0 2 -2 0 2 0T1 0 1 -1 0 2 N=13 -20/13 -21/13 MPI -1.54 -1.62

Malvaceae Asteraceae Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) BR1TOP 1 2 1 -1 2 BR1MID 0 1 0 4 BR1 BASE 1 0 -1 2 BR7 TOP 1 0 -1 3 BR7MID 1 0 -1 4 BR1 BASE 1 0 -1 2 BR2 0 2 2 -1 2 BR3 TOP 1 2 1 -1 3 -338-

BR3MK) 2 1 0 2 BR3 BASE 2 1 0 2 BR4 (10cm) 1 0 0 2 BR4 (20cm) 1 0 0 2 BR4 (30cm) 2 2 -1 2 BR4 (40cm) 1 0 0 2 BR4 (50cm) 1 0 0 3 BR4 (60cm) 1 0 0 3 N= 16 -9/16 -32/16 MPI -0.56 -2

Apiaceae Myoporaceae Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) BR1 TOP 1 2 -1 1 2 BR1MID 0 2 -2 0 2 BR1 BASE 0 2 -2 0 3 BR7 TOP 0 2 -2 0 2 BR7MID 0 2 -2 0 2 BR1 BASE 0 2 -2 0 2 BR2 0 1 -1 0 2 BR3 TOP 11 2 -1 0 2 BR3MID 1 2 -1 0 2 BR3 BASE 0 2 -2 0 2 BR4 (10cm) 1 2 -1 0 2 BR4 (20cm) 1 2 -1 1 2 BR4 (30cm) 1 3 -2 0 2 BR4 (40cm) 1 2 -1 0 2 BR4 (50cm) 0 2 -2 1 2 BR4 (60cm) 0 2 -2 0 2 N= 16 -25/16 -30/16 MPI -1.56 -1.06

Acacia Sapindaceae Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) BR1 TOP 0 1 -1 1 1 BR1MID 1 1 0 0 2 BR1 BASE 1 1 0 1 1 BR7 TOP 0 2 -2 1 2 -339-

BR7MID 0 1 -1 1 2 BR1 BASE 0 2 -2 1 2 BR2 0 2 -2 0 2 BR3 TOP 1 2 -1 0 1 BR3MID 1 2 -1 1 2 BR3 BASE 1 2 -1 0 2 BR4 (10cm) 1 2 -1 0 2 BR4 (20cm) 1 2 -1 1 2 BR4 (30cm) 1 2 -1 1 3 BR4 (40cm) 1 1 0 1 2 BR4 (50cm) 0 2 -2 1 2 BR4 (60cm) 0 1 -1 0 2 N=16 -17/16 -20/16 MPI -1.06 -1.25

Casuarinaceae Poaceae Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen BR1 TOP 0 1 -1 0 2 BR1MID 0 1 -1 0 2 BR1 BASE 0 2 -2 0 2 BR7 TOP 0 2 -2 0 2 BR7MID 0 2 -2 0 2 BR1 BASE 0 2 -2 1 2 BR2 2 -1 0 2 BR3 TOP 2 -1 0 2 BR3MID 2 -1 1 2 BR3 BASE 2 -1 0 2 BR4 (10cm) 2 -1 0 2 BR4 (20cm) 2 -1 0 2 BR4 (30cm) 0 2 -2 0 2 BR4 (40cm) 0 2 -2 0 2 BR4 (50cm) 0 2 -2 0 2 BR4 (60cm) 0 1 -1 0 2 N= 16 -23/16 -30/16 MPI -1.44 -1.88 -340-

Chenopodiaceae/ Amaranthaceae Midden Code Macrofossil (M) Pollen (P) M-P BR1TOP 0 4 -4 BR1MID 0 4 -4 BR1 BASE 0 3 -3 BR7 TOP 1 3 -2 BR7MID 0 4 -4 BR1 BASE 0 3 -3 BR2 0 5 -5 BR3 TOP 1 5 -4 BR3MID 1 5 -4 BR3 BASE 0 4 -4 BR4 (10cm) 1 4 -3 BR4 (20cm) 1 4 -3 BR4 (30cm) 0 4 -4 BR4 (40cm) 1 5 -4 BR4 (50cm) 0 4 -4 BR4 (60cm) 0 4 -4 N= 16 -59/16 MPI -3.69

Malvaceae Poaceae Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) M- MCI EXP 1 1 0 0 1 -1 MCI TOP 1 1 0 0 1 -1 MC2TOP 1 1 0 1 1 0 MC2 OVERHG 1 2 -1 0 2 -2 CG 0 1 -1 1 1 0 MD3 1 2 -1 0 2 -2 N=6 -3/6 -6/6 MPI -0.5 -1

Chenopodiaceae/ Amaranthaceae Myoporaceae Midden Code Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) M MCI EXP 0 5-5 0 2-2 MCI TOP 0 5-5 0 2-2 MC2TOP 1 5 -4 1 2 -1 MC2 OVERHG 1 4-3 0 2-2 -341-

CG 0 3 -2 0 2 -2 MD3 0 4 -4 0 2 -2 N=6 -23/6 -11/6 MPI -3.8 -1.83

Casuarinaceae Poaceae Middens (7-5ka) Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) M BR1 TOP 0 1 -1 0 2 -2 BR1MID 0 1 -1 0 2 -2 BR1 BASE 0 2 -2 0 2 -2 BR2 1 2 -1 0 2 -2 BR7 TOP 0 2 -2 0 2 -2 BR7MID 0 2 -2 0 2 -2 BR7 BASE 0 2 -2 1 2 -1 BR4 (10cm) 1 2 -1 0 2 -2 BR4 (20cm) 1 2 -1 0 2 -2 BR4 (30cm) 0 2 -2 0 2 -2 BR4 (40cm) 0 2 -2 0 2 -2 BR4 (50cm) 0 2 -2 0 2 -2 BR4 (60cm) 0 1 -1 0 2 -2 MD3 0 0 0 0 2 -2 N= 14 -20/14 -27/14 MPI -1.43 -1.93

Chenopodiaceae/ Amaranthaceae Malvaceae Middens (7-5ka) Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) M BR1 TOP 0 4 -4 1 2 -2 BR1MID 0 4 -4 0 -1 BR1 BASE 0 3 -3 1 0 BR2 0 5 -5 0 2 -2 BR7 TOP 1 3 -2 1 0 BR7MID 0 4 -4 1 0 BR7 BASE 0 3 -3 1 0 BR4 (10cm) 1 4 -3 1 0 BR4 (20cm) 1 4 -3 1 0 BR4 (30cm) 0 4 -4 1 2 -1 BR4 (40cm) 1 5 -4 1 0 BR4 (50cm) 0 4 -4 1 0 -342-

BR4 (60cm) 0 4 -4 1 1 MD3 0 4 -4 1 2 N= 14 -51/14 -7/14 MPI -3.64 -0.5

Sapindaceae Asteraceae Middens (7-5ka) Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) BR1 TOP 1 1 0 1 2 BR1MID 0 2 -2 0 4 BR1 BASE 1 1 0 2 BR2 0 2 -2 2 BR7 TOP 1 2 -1 3 BR7MED 1 2 -1 4 BR7 BASE 1 2 -1 2 BR4 (10cm) 0 2 -2 0 2 BR4 (20cm) 1 2 -1 0 2 BR4 (30cm) 1 3 -2 1 2 BR4 (40cm) 1 2 -1 0 2 BR4 (50cm) 1 2 -1 0 2 BR4 (60cm) 0 2 -2 0 3 MD3 0 2 -2 0 2 N= 14 -18/14 -30/14 MPI -1.29 -2.14

Apiaceae Myoporaceae Middens (7-5ka) Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) BR1 TOP 1 2 -1 1 2 BR1MID 0 2 -2 0 2 BR1 BASE 0 2 -2 0 3 BR2 0 1 -1 0 2 BR7 TOP 0 2 -2 0 2 BR7MID 0 2 -2 0 2 BR7 BASE 0 2 -2 0 2 BR4 (10cm) 1 2 -1 0 2 BR4 (20cm) 1 2 -1 1 2 BR4 (30cm) 1 3 -2 0 2 BR4 (40cm) 1 2 -1 0 2 BR4 (50cm) 0 2 -2 1 2 -343-

BR4 (60cm) 0 2 -2 0 2 -2 MD3 0 2 -2 0 2 -2 N= 14 -23/14 -26/14 MPI -1.64 -1.86

Acacia Middens (7-5ka) Macrofossil (M) Pollen (P) M-P BR1TOP 0 1 -1 BR1MID 1 1 0 BR1 BASE 1 1 0 BR2 0 2 -2 BR7 TOP 0 2 -2 BR7MID 0 1 -1 BR7 BASE 0 2 -2 BR4 (10cm) 1 2 -1 BR4 (20cm) 1 2 -1 BR4 (30cm) 1 2 -1 BR4 (40cm) 1 1 0 BR4 (50cm) 0 2 -2 BR4 (60cm) 0 1 -1 MD3 0 2 -2 N= 14 -16/14 MPI -1.14

Malvaceae Asteraceae Middens (4-2ka) Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) M-P BR3 TOP 2 -1 1 3 -2 BR3MID 2 -1 0 2 -2 BR3 BASE 2 -1 0 2 -2 NWCK 2 1 0 0 4 -4 RC1 2 -1 0 3 -3 RC2 0 1 -1 1 3 -2 MPI -0.83 -2.5 -344-

Apiaceae Acacia Middens (4-2ka) Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) BR3 TOP 1 2 -1 1 2 BR3MID 1 2 -1 1 2 BR3 BASE 0 2 -2 1 2 NWCK 2 0 2 -2 1 1 RC1 1 2 -1 0 1 RC2 1 2 -1 1 2 MPI -1.33 -0.83

Sapindaceae Casuarinaceae Middens (4-2ka) Macrofossil (M) Pollen (P) M-P Macrofossil (M) Pollen (P) BR3 TOP 0 1 -1 1 2 BR3MID 1 2 -1 1 2 BR3 BASE 0 2 -2 1 2 NWCK 2 0 2 -2 0 2 RC1 0 2 -2 1 1 RC2 0 2 -2 0 2 MPI -1.67 -1.17

Chenopodiaceae/ Amaranthaceae Poaceae Middens (4-2ka Macrofossil (M) Pollen (P) M-4P Macrofossil (M) Pollen (P) BR3 TOP 1 5 -4 0 2 BR3MID 1 5 -4 1 2 BR3 BASE 0 4 -4 0 2 NWCK 2 0 3 -3 0 2 RC1 1 2 -1 1 2 RC2 1 3 -2 1 2 MPI -3 -1.5

Malvaceae Poaceae Middens (

ARK1BASE 1 1 0 0 2 HHl TOP 0 1 -1 0 2 HHl BASE 0 2 -2 0 2 0T1 0 1 -1 0 1 MCI EXP 1 1 0 0 1 MCI TOP 1 1 0 0 1 MC2TOP 1 1 0 1 1 MC2 OVERHG 1 2 -1 0 2 CGI 0 1 -1 1 1 N= 15 -11/15 -21/15 MPI -0.73 -1.4

Chenopodiaceae/ Amaranthaceae Myoporaceae Middens (

Asteraceae Casuarinaceae Middens (

ARK1 TOP 1 4 3 0 -1 ARK1MID 1 3 2 0 -1 ARK1 BASE 2 5 3 0 -1 HHl TOP 0 3 3 0 -2 HHl BASE 0 3 3 0 -1 0T1 0 2 2 1 0 MCI EXP 0 2 2 0 -1 MCI TOP 0 2 2 0 -1 MC2 TOP 0 2 2 0 -1 MC2 OVERHG 0 4 4 0 0 0 CGI 0 2 2 0 1 -1 N= 15 -37/15 -15/15 MPI -2.47 -1

Apiaceae Acacia Middens (

Sapindaceae Caesalpinaceae Middens (

Sorenson Similarity Index (SI) Values for Compariso Between Modern Pollen Rain and Modern Vegetation

SI = 2C / A + B where A= number of pollen taxa in the modern pollen rain B= number of pollen taxa recorded in the vegetation C= number of common taxa

A) West-east Transect Modern Pollen Study

Site and Trap No. Calculation SI Arkaroola 1995 Trapl 2x5/25+5 0.33 Trap 10 2x3/19+3 0.27 Trap 11 2x5/20+5 0.40 Trap 23 2x5/21+6 0.37 Chambers Gorge 1995 Trap 9 2x1/22+2 0.08 Trap 11 2x2/17+3 0.2 Trap 13 2x2/18+2 0.2 Brachina Gorge 1995 Trap 8 2x2/19+3 0.18 Trap 9 2x2/22+2 0.17

B) Midden Cave Site Modern Pollen Study

Site and Trap No. Calculation SI Arkaroola 1995 Trap 2 2x7/16+11 0.52 Trap 3 2x9/20+11 0.58 Trap 6 2x5/21+10 0.32 Trap 7 2x6/21+10 0.39 Trap 4 2x8 / 23+10 0.48 Trap 5 2x7/15+10 0.56 Trap 9 2x5 /17+6 0.43 Trap 19 2x9/19+12 0.58 Trap 20 2x11/26+12 0.57 Trap 17 2x8/17+13 0.53 Trap 18 2x8 / 15+13 0.57 Trap 15 2x3 / 20+4 0 Trap 16 2x4 / 23+4 0 Trap 21 2x8 / 19+10 0 Trap 22 2x4 / 8+10 0 Chambers Gorge 1995 Trap 1 2x4 / 24+5 0 Trap 4 2x7 / 23+9 0 Trap 3 2x6 / 20+9 0 Trap 5 2x5 / 12+7 0 Trap 7 2x7 / 17+8 0 Trap 8 2x7 / 22+8 0 Brachina Gorge 1995 Trap 10 2x12/26+14 0 Trapl 2x8/21+14 0 Trap 5 2x10/24+13 0 Trap 6 2x10/24+13 0 -350-

APPENDIX 10: PLANT LIST FOR SPECIES RECORDED AT MIDDEN SITES IN THE NORTHERN AND CENTRAL FLINDERS RANGES

Plant Species Recorded at Midden Sites Arkaroola- Brachina Mount in the central and northern Flinders Mount Gorge Chambers Ranges Painter Gorge Sanctuary Abutilon leucopetalum * * * Acacia aneura * * Acacia confluens * Acacia continua * Acacia ligulata * Acacia rivalis * * Acacia salicina * Acacia tetragonophylla * * * Acacia victoriae * * * Acetosa vesicarius * * * Alectryon oleifolium * * * Amymea miquelii * Anagallis arvensis * * Arabidellafilfolia * Aristida nitidula * Arthropodium strictum * Avena barbata * Brachycome ciliaris spp. ciliaris * Brachycome ciliaris var lyrifolia * Brachycome spp. * Bulbine semibarbatus * Bursaria spinosa * * * Callitris columellaris * * * Calotis latiuscula * Capparis mitchelli * Carrichtera annua * Cassinia laevis * * * Casuarina cristata * * * -351-

Casuarina pauper * Casuarina verticulata * Cheilanthes austrotenuifolia * *

Cheilanthes lasiophylla * Cheilanthes sieberi * Chenopodium cristatum * Chenopodium melanocarpum * Chenopodium spp. * * Chrysocephalem semicaluum * Chrysocephalem semicaluum spp. * * semicaluum Chrysocephalem semipapposum * Citrillus lanatus * * Clianthus formosus * Convolvulus remotus * Cymbopogan ambiguus * * * Cyperaceae gillesii * * Cyperaceae spp. * Danthonia aspitosa * Danthonia tenuior * Daviesia genistifolia * Dicondra repens * * Digitaria brownii * Dissocarpus paradoxus * * Dodonaea lobulata * Dodonaea microzya * Dodonaea viscosa * Dodonaea viscosa spp. angustissima * * * Dodonaea viscosa spp. spatulata * Echium plantagenium * * Enchylaena tomentosa * * * Enneapogan arenaceus * Enneapogon avenaceus {check spelling) * Enneapogon polyphyllus * * Eremophila alternifolia * * Eremophila freelingii -352-

Eremophila latrobei * Eremophila longifolia # Eremophila scopana * Eriachne mucronata * Eucalyptus camaldulensis * * * Eucalyptus camaldulensis var. obtusa * Eucalyptus intertexta * Eucalyptus socialis * Euphorbia drummondii * * *

Exocarpos aphyllus * * * Galium migrans * Galium spp. * Glycine clandestina * Goodenia ovata * * Goodenia vernicosa * * Hakea ednieana * * * Hakea leucoptera * Indigofera leucotricha * Isotoma petraea * Ixolaena leptolepis * Lepidium papillosum * Lomandra multiflora ssp. dura * Lotus cruentus * Lysiana exocarpi * * Maireana brevifolia * Maireana eriodada or pentatropis * Maireana georgei * Maireana pyramidata * * Marrubium vulgare * * Melalecua glomerata * * * Melaleuca dissitiflora * Meliotus indica * Montanum desertii * Mvoporum montanum * Myoporum platscarpum * * * Nicotiana glauca -353-

Olearia decurrens * * * Olearia pimeleoides *

Olearia stuartii * * Paraceterach reynoldsii *

Paspalidium constrictum * ! Pimelea microcephala * * * i Pimelea microphylla * Pluchea dentex * * Polypogan maritimus * * Portulaca oleracea * Prostanthera striatiflora * * * Pterocaulon sphacelatum * * Ptilotus exultatus * Ptilotus obovatus * * * Ranunculus hamatosetosus * Rhagodia parabolica * Salso kali * Santalum lanceolatum * Scaevola spinescens * * Schoenoplectus littoralis * Sclerolaena cuneata * Sclerolaena holtiana * Senecio magnificus * * Senna artemisiodies spp. helmsii * Senna artemisiodies x ssp. sturtii * Senna artemisioides ssp. artemisioides * * * Senna artemisioides ssp. artemisioides * (affinities to petiolaris) Senna artemisioides x ssp. coriacea * Senna oligophylla * Sida fibulifera * * * Sida petrophila * Sigesbeckia microcephala * * Sisymbrium erysimoides * * * * * Solanum ellipticum * * Solanum petrophilum * * -354-

Stipa eremophila * Stipa scalra spp.taleata * Stipa scalra ssp. falcata * Stipa spp. * Tetragonia eremaea * Tetragonia tetragonioides * Themeda triandra * Trachymene glaucifolia * *

Trichodesma zeylanicum * * Triodia irritans * * Typha domingensis * * Urospermum picroides * Vittadinia cuneata * Xanthorrhoea quadrangulata * * Zygophyllum apiculatum * * * Zygophyllum aurantiacum * *