Agricultural Development in Tongan Prehistory: an Archaeobotanical Perspective

AGRICULTURAL DEVELOPMENT IN TONGAN PREHISTORY: AN ARCHAEOBOTANICAL PERSPECTIVE

A Dissertation by ELLA USSHER

A thesis submitted for the degree of Doctor of Philosophy

at the Australian National University

June 2015

School of Culture, History and Language

Department of Archaeology and Natural History

DECLARATION

The research presented here is based on original fieldwork, as well as analysis of micro- and macrobotanical assemblages excavated by the author on Tongatapu, Kingdom of Tonga. I certify that, except where it is stated otherwise, this dissertation is the result of my own original investigation.

Ella Ussher

Acknowledgements

The completion of this thesis would not have been possible without the contribution and support of a number of people. Firstly, I would like to thank my Chair of Panel Assoc. Prof. Geoff Clark for proposing this project and supporting my fieldwork in Tonga, as well as guiding the final shaping of the written component of my research. My other supervisors, especially Janelle Stevenson and Matiu Prebble, provided me with invaluable technical support over the last four years in both the construction of my comparative collection and the analysis of archaeological material. Many other staff and students at the ANU provided me with specialised advice. These included Dr Jack Fenner (statistics), Dr Frank Brinks (SEM), Dr Anne Prins (histology), Feli Hopf and Jay Chin (microbotanical labwork), Rose Whitau (macrobotanical labwork) and Maxine MacArthur (copy editing). To these people, I am very grateful for their time and sharing of experience and equipment, without which this research could not have been completed.

On a more personal level, I wish to give a massive thank you to my partner Josh for being so understanding of the time and effort that needed to go into this PhD project. His love and support ensured that I retained my sanity at the very end and was well looked after as we prepared to welcome our little girl into the world. Thanks and lots of love also needs to go to my family back in NZ who encouraged me to go to Canberra and take my academic career further so that I could follow my passion for archaeobotany. Close friends and colleagues from the ANU such as Katherine Seikel, Stuart Hawkins, Jay Chin, Rebecca Jones, Mirani Litster, Feli Hopf, Christian Reepmeyer, Tim Maloney, Rose Whitau, Ben Shaw, Justin Lewis, and Matthieu Leclerc all helped create a fun and collegial atmosphere both on and off campus. Finally, I would also like to express my gratitude to those who helped distract me from my thesis when I really needed a break by allowing me to follow my other passion in horses, and providing amazing friendships along the way that I will value forever (Maxine, Ann, Wendy, Keryn, Claire, Cathy, Jeremy, Fia, and Kaaren).

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Abstract

This thesis presents the results of an archaeobotanical study of agricultural development in the Kingdom of Tonga. Prior to this study, there has been no direct archaeological evidence for agriculture in Tongan prehistory. Through the implementation of systematic archaeobotanical techniques, this study aimed to fill this gap and address two key research questions: 1) whether early colonisers were dependent on introduced crops, or if human dispersal was fuelled predominantly by the exploitation of natural resources; and 2) whether archaeobotanical data can provide new evidence to examine the role of agriculture within the development of the maritime chiefdom in Tonga through agroecological modelling.

This research was divided into two main phases. The first involved the construction of a comprehensive comparative collection for macrobotanical (vegetative storage and fruit parenchyma and endocarp), and microbotanical (starch) components of economic and supplementary plant taxa from Tonga. As part of this, a study of the morphological attributes of starch and parenchyma was conducted that incorporated multivariate statistical analyses of diagnostic attributes. Two methods for taxonomic classification were suggested: automated classification using Discriminant Function Analysis (DFA) of starch, and the use of an Identification Flowchart Key for parenchyma.

In the second phase of research, archaeobotanical data from three sites on Tongatapu, representing three different time periods in Tongan prehistory, is presented. Macrobotanical and microbotanical remains were extracted from these sites using flotation, wet-sieving and bulk stratigraphic sampling and compared to a comprehensive reference collection using a combination of SEM and light microscopy. Sampled cultural deposits at Talasiu (2750-2650 cal BP), Leka (1300-1000 cal BP) and Heketa (800-600 cal BP) present new insights into the role of plant taxa within late-Lapita, the Formative Period, and early stages of the Classic Tu’i Tonga chiefdom. Modelling using techniques from Human Ecology, specifically agroecology, replicated past production systems using measures of system efficiency such as nutritional value of taxa, labour investment and productivity in terms of yields. These were compared to expectations based on current literature, and a revised chronology for agricultural development and links to social complexity is presented.

This study demonstrates that multivariate statistical analysis and identification flowcharts enable the discrimination of starch and vegetative storage parenchyma from most Tongan plant taxa based on metric and nominal morphological attributes. When applied to archaeobotanical data these techniques indicate that most staple cultigens and some supplementary or famine foods were brought to Tonga within a few hundred years of initial Lapita colonisation. Late prehistoric introductions likely included the sweet potato (Ipomoea

iv batatas) by 600 BP, transported via East Polynesia through the extensive trade networks of the developing Tongan state. Modelling past production systems linked decreased system nutritional efficiency over time to horticultural specialisation in primary crops and increasingly centralised government on Tongatapu. Critically, this analysis modelled the high nutritional efficiency of Lapita subsistence, and linked this to the division of labour investment between both economic and supplementary species within a decentralised social hierarchy.

Keywords: archaeobotany, starch, parenchyma, microfossils, Tonga, archaeology, agriculture, agroecology, production systems, Lapita

Table of Contents

Acknowledgements iii Abstract iv Table of Contents vi List of Tables ix List of Figures xiii List of Abbreviations xvii Chapter 1 Introduction 1 Research aims and objectives 1 Theoretical framework 5 Thesis organisation 6

Chapter 2 Tongan Agriculture in the Pacific Context 9 Research to date 9 Geographic and climatic limitations to agricultural modelling 10 Ethno-historic accounts of plant cultivation in Tonga: 12 Archaeobotany of cultigens in the Pacific 20 Tonga in the Pacific: A summary 30

PART ONE- AN ARCHAEOBOTANICAL COMPARATIVE COLLECTION FOR TONGA 32 Chapter 3 Reviewing Microbotanical Analysis 33 Biology of starch and identification potential 33 Starch taphonomy 35 Modern starch contamination 42 Sampling strategies and extraction techniques 44

Chapter 4 Reviewing Parenchyma 48 Fresh and charred parenchyma morphology 48 Taphonomic factors affecting macrobotanical preservation 49 Collection and sampling of parenchyma 52 Parenchyma identification 54

Chapter 5 Comparative Collection and Morphometric Studies of Pacific Cultigens 57 Species selection 57 Field collection 58 Laboratory processing of samples 59 Starch processing 59 Histology 60 Experimental charring 61 Recording 62 Light microscopy 62 Scanning Electron Microscopy 62 Morphology of native starch 63 Starch morphology 64 Multivariate statistical analysis of starch 76 Morphology of vegetative storage parenchyma 82 Morphological analysis of fresh samples 82 Description of charcoal 100 Development of an Identification Flowchart Key 106

PART TWO- DEVELOPMENT OF PREHISTORIC AGRICULTURE IN TONGA 112 Chapter 6 Sites and Field Sampling Strategy 113 Methodology for field sampling of archaeological sediments 113 Site selection 113 Field methods 114 Site descriptions 116 Talasiu (TO-Mu-2) 116 Leka (J17) 117 Heketa (TO-Nt-2) 118 Stratigraphic descriptions 119 Talasiu (TO-Mu-2) 119 Leka (J17) 120 Heketa (TO-Nt-2) 122 AMS dating of cultural contexts 124

Chapter 7 Laboratory Methods 126 Microbotanical analysis: Starch residues 126 Experimentation with starch extraction techniques 126 Laboratory processing: Revised starch extraction protocol 132 Light Microscopy 134 Archaeological starch classification: Assemblage-typology approach 134 Archaeological starch classification: Multivariate statistical analysis 135 Macrobotanical analysis: Charred parenchyma and endocarp 136 Laboratory analysis 136

Chapter 8 Results 138 Macrobotanical analysis 138 Quantification of charcoal 138 Parenchyma distribution and identification: Talasiu TP2 case study 142 Microbotanical analysis 144 Extraction, quantification and distribution 144 Identification: Assemblage-typology approach 147 Identification: Multivariate statistics—Discriminant Function Analysis 150 Comparison of modern Pacific production systems 160 Nutrition 161 Labour investment 178 Outputs 187 Output to input ratios: Efficiency calculation 194 System efficiency comparison and system classification 202 Comparison of prehistoric production systems 209 Nutritional comparison of archaeological species 209 Efficiency comparison of archaeological species and production systems 215

Chapter 9 Discussion 221 Timing and nature of plant introductions into Tonga 221 Spondias dulcis— Anacardiaceae 222 Alocasia macrorrhiza— Araceae 222 Amorphophallus paeoniifolius— Araceae 223 Colocasia esculenta— Araceae 223 Cyrtosperma merkusii— Araceae 224 Cocos nucifera— Arecaceae 225 Ipomoea batatas— Convolvulaceae 225 vii

Dioscorea spp.— Dioscoreaceae 226 Inocarpus fagifer— Fabaceae 227 Barringtonia asiatica— Lechythidaceae 228 Artocarpus altilis— Moraceae 228 Musa spp.— Musaceae 229 Piper methysticum— Piperaceae 230 Curcuma longa and Zingiber spp.— Zingiberaceae 231 Modelling archaeological production systems 232 Feasibility of modelling 232 Expected modelling outcomes 234 Modelling Talasiu (TO-Mu-2) 240 Modelling Leka (J17) 244 Modelling Heketa (TO-Nt-2) 247 Comparison of expected and modelled outcomes 250 Specialisation and system efficiency 255 Contamination at Leka and Heketa 256 Linking archaeobotanical data to island colonisation and social complexity 257

Chapter 10 Conclusion 260 Meeting research aims and objectives 260 Future recommendations 267 Micro- and macrobotanical techniques 267 Archaeobotanical research in Tonga and the Pacific 269

Bibliography 271 Appendix A- Species in Reference Collection 294 Appendix B- Description of Parenchyma 296 Appendix C- Starch Images 326 Appendix D- Archaeobotanical Research in the Pacific 331

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List of Tables

TABLE 2.1 LIST OF SPECIES RECORDED IN EARLY ETHNO-HISTORIC ACCOUNTS FROM TONGA...... 19 TABLE 5.1 HILUM FISSURING OF REFERENCE SPECIES ...... 66 TABLE 5.2 THREE-DIMENSIONAL SHAPES OF REFERENCE SPECIES ...... 68 TABLE 5.3 SUMMARY OF STARCH MORPHOLOGY WITHIN REFERENCE COLLECTION ...... 74 TABLE 5.4 DESCRIPTION OF METRIC AND BINARY VARIABLES USED DURING DISCRIMINANT FUNCTION ANALYSIS...... 76 TABLE 5.5 SPECIES INCLUDED IN CENTRIC AND ECCENTRIC DATASETS FOR TONGAN ANALYSIS ...... 78 TABLE 5.6 GROUND TISSUE CELL SHAPES OF TAXA IN REFERENCE COLLECTION ...... 84 TABLE 5.7 GROUND TISSUE CELL DIMENSIONS OF TAXA IN THE REFERENCE COLLECTION ...... 84 TABLE 5.8 VASCULAR TISSUE ARRANGEMENTS OF TAXA IN REFERENCE COLLECTION ...... 91 TABLE 5.9 SUMMARY OF PARENCHYMA MORPHOLOGY WITHIN REFERENCE COLLECTION ...... 96 TABLE 5.10 DESCRIPTION OF MORPHOLOGICAL MODIFICATION WITHIN GROUND TISSUE OF CHARRED SAMPLES IN THE COMPARATIVE COLLECTION ...... 103 TABLE 5.11 DESCRIPTION OF MORPHOLOGICAL MODIFICATION WITHIN VASCULAR TISSUE OF CHARRED SAMPLES IN THE COMPARATIVE COLLECTION ...... 104 TABLE 8.1 SUMMARY OF TOTAL MACROBOTANICAL ASSEMBLAGES FROM ALL SITES AND TEST-PITS ...... 139 TABLE 8.2 QUANTIFICATION OF COCONUT ENDOCARP FROM ALL SITES AND TEST-PITS ...... 140 TABLE 8.3 QUANTIFICATION OF OTHER ENDOCARP FROM ALL TEST UNITS ...... 141 TABLE 8.4 QUANTIFICATION OF WOOD CHARCOAL AND PARENCHYMA FROM ALL TEST UNITS ...... 142 TABLE 8.5 DISTRIBUTION AND IDENTIFICATION OF PARENCHYMA EXTRACTED FROM TALASIU TP2 ...... 144 TABLE 8.6 OVERALL QUANTITIES (COUNTS) OF STARCH EXTRACTED FROM ALL SAMPLED TEST UNITS AT TALASIU (TO-MU-2) ...... 145 TABLE 8.7 DISTRIBUTION OF STARCH COUNTS WITHIN TALASIU TP2 ...... 146 TABLE 8.8 DISTRIBUTION OF STARCH COUNTS WITHIN LEKA TP2 ...... 146 TABLE 8.9 DISTRIBUTION OF STARCH COUNTS WITHIN LEKA TP4 ...... 147 TABLE 8.10 DISTRIBUTION OF STARCH COUNTS WITHIN HEKETA TP3 ...... 147 TABLE 8.11 TABLE OUTLINING SUGGESTED FAMILY OF ORIGIN OF ARCHAEOLOGICAL STARCH TYPES FROM TALASIU TP2...... 149 TABLE 8.12 DISTRIBUTION OF PRELIMINARY IDENTIFICATIONS WITHIN TALASIU TP2 USING THE ASSEMBLAGE-TYPOLOGY APPROACH ...... 150 TABLE 8.13 LEVELS OF CONFIDENCE FROM DFA CLASSIFICATION OF ARCHAEOLOGICAL STARCH FROM

TALASIU TP2. NB HIGH CONFIDENCE (BLACK), MODERATED CONFIDENCE (MEDIUM GREY) AND LOW CONFIDENCE (LIGHT GREY) ...... 153 TABLE 8.14 FINAL TABLE DOCUMENTING SPECIES REPRESENTED BY ARCHAEOLOGICAL STARCH WITHIN TALASIU TP2 NB PRESENCE INDICATED BY BLACK SQUARES ...... 155 TABLE 8.15 LEVELS OF CONFIDENCE FROM DFA CLASSIFICATION OF ARCHAEOLOGICAL STARCH FROM

LEKA TP2. NB HIGH CONFIDENCE (BLACK), MODERATED CONFIDENCE (MEDIUM GREY) AND LOW CONFIDENCE (LIGHT GREY) ...... 155 TABLE 8.16 FINAL TABLE DOCUMENTING SPECIES REPRESENTED BY ARCHAEOLOGICAL STARCH WITHIN LEKA TP2 ...... 156 ix

TABLE 8.17 LEVELS OF CONFIDENCE FROM DFA CLASSIFICATION OF ARCHAEOLOGICAL STARCH FROM

LEKA TP4. NB HIGH CONFIDENCE (BLACK), MODERATED CONFIDENCE (MEDIUM GREY) AND LOW CONFIDENCE (LIGHT GREY) ...... 156 TABLE 8.18 FINAL TABLE DOCUMENTING SPECIES REPRESENTED BY ARCHAEOLOGICAL STARCH WITHIN LEKA TP4 ...... 157 TABLE 8.19 LEVELS OF CONFIDENCE FROM DFA CLASSIFICATION OF ARCHAEOLOGICAL STARCH FROM

HEKETA TP3. NB HIGH CONFIDENCE (BLACK), MODERATED CONFIDENCE (MEDIUM GREY) AND LOW CONFIDENCE (LIGHT GREY) ...... 158 TABLE 8.20 FINAL TABLE DOCUMENTING SPECIES REPRESENTED BY ARCHAEOLOGICAL STARCH WITHIN HEKETA TP3 ...... 158 TABLE 8.21 NUTRITIONAL FIGURES AND RANKINGS FOR SPECIES WITHIN THE GADIO ENGA SYSTEM

ACCORDING TO CALORIES, PROTEIN, FATS, CARBOHYDRATES AND TOTAL NUTRITION FIGURES (DATA FROM DORNSTREICH 1974, 1978) ...... 163 TABLE 8.22 NUTRITIONAL FIGURES AND RANKINGS FOR SPECIES WITHIN BELLONA ISLAND SYSTEM

ACCORDING TO CALORIES, PROTEIN, FATS, CARBOHYDRATES AND TOTAL NUTRITION FIGURES (DATA FROM CHRISTIANSEN 1975) ...... 166 TABLE 8.23 NUTRITIONAL FIGURES AND RANKINGS FOR SPECIES WITHIN ANUTAN SYSTEM ACCORDING TO

CALORIES, PROTEIN, FATS, CARBOHYDRATES AND TOTAL NUTRITION FIGURES (DATA FROM YEN 1973B) ...... 170 TABLE 8.24 NUTRITIONAL FIGURES AND RANKINGS FOR SPECIES WITHIN TONGAN SYSTEM ACCORDING TO

CALORIES, PROTEIN, FATS, CARBOHYDRATES AND TOTAL NUTRITION FIGURES (DATA FROM MINISTRY OF AGRICULTURE AND FORESTRY 2001) ...... 173 TABLE 8.25 NUTRITIONAL FIGURES AND RANKINGS FOR SPECIES WITHIN ONTONG JAVA SYSTEM

ACCORDING TO CALORIES, PROTEIN, FATS, CARBOHYDRATES AND TOTAL NUTRITION FIGURES SYSTEM (DATA FROM BAYLISS-SMITH 1973, 1986) ...... 176 TABLE 8.26 STATISTICAL COMPARISON OF SPECIES’ GROUPINGS IN EXAMPLE SYSTEMS ACCORDING TO OVERALL NUTRITION FIGURES/100G ...... 178 TABLE 8.27 LABOUR INVESTMENT INTO SPECIES WITHIN THE GADIO ENGA SYSTEM (DATA FROM DORNSTREICH 1974, 1978) ...... 179 TABLE 8.28 LABOUR INVESTMENT INTO SPECIES WITHIN THE BELLONA IS SYSTEM (DATA FROM CHRISTIANSEN 1975) ...... 181 TABLE 8.29 LABOUR INVESTMENT INTO SPECIES WITHIN THE ANUTAN SYSTEM (DATA FROM YEN 1973B) ...... 182 TABLE 8.30 LABOUR INVESTMENT INTO SPECIES WITHIN THE TONGAN SYSTEM (DATA FROM MINISTRY OF AGRICULTURE AND FORESTRY 2001) ...... 184 TABLE 8.31 LABOUR INVESTMENT INTO SPECIES WITHIN THE ONTONG JAVA PLANT PRODUCTION SYSTEM (DATA FROM BAYLISS-SMITH 1973, 1986) ...... 185 TABLE 8.32 STATISTICAL COMPARISON OF SPECIES’ GROUPINGS IN EXAMPLE SYSTEMS ACCORDING TO LABOUR INPUT FIGURES ...... 187 TABLE 8.33 OUTPUT COMPARISON OF SPECIES IN GADIO ENGA PLANT PRODUCTION SYSTEM (DATA FROM DORNSTREICH 1974, 1978) ...... 188

TABLE 8.34 OUTPUT COMPARISON OF SPECIES IN BELLONA ISLAND SYSTEM (DATA FROM CHRISTIANSEN 1975) ...... 189 TABLE 8.35 OUTPUT COMPARISON OF SPECIES IN ANUTAN SYSTEM (DATA FROM YEN 1973B)...... 191 TABLE 8.36 OUTPUT COMPARISON OF SPECIES IN TONGAN SYSTEM (DATA FROM MINISTRY OF AGRICULTURE AND FORESTRY 2001) ...... 192 TABLE 8.37 OUTPUT COMPARISON OF SPECIES IN ONTONG JAVA PRODUCTION SYSTEM (DATA FROM BAYLISS-SMITH 1973, 1986) ...... 194 TABLE 8.38 YIELD RATIOS FOR ARCHAEOLOGICAL SPECIES IN ALL MODERN PRODUCTION SYSTEMS (KG/TIME UNIT OF LABOUR) ...... 196 TABLE 8.39 OUTPUT TO INPUT RATIOS FOR ARCHAEOLOGICAL SPECIES USING GADIO ENGA DATA ...... 198 TABLE 8.40 OUTPUT TO INPUT RATIOS FOR ARCHAEOLOGICAL SPECIES USING BELLONA DATA ...... 199 TABLE 8.41 OUTPUT TO INPUT RATIOS FOR ARCHAEOLOGICAL SPECIES USING ANUTAN DATA ...... 200 TABLE 8.42 OUTPUT TO INPUT RATIOS FOR ARCHAEOLOGICAL SPECIES USING TONGAN 2001 DATA ...... 201 TABLE 8.43 OUTPUT TO INPUT RATIOS FOR ARCHAEOLOGICAL SPECIES USING ONTONG JAVAN DATA ..... 202 TABLE 8.44 STATISTICAL COMPARISON OF NUTRITIONAL VALUE OF ARCHAEOLOGICAL AND EXPECTED ETHNOGRAPHIC SPECIES ...... 215 TABLE 8.45 STATISTICAL COMPARISON OF NUTRITIONAL VALUE OF SPECIES GROUPS WITHIN ARCHAEOLOGICAL SYSTEMS AT TALASIU, LEKA AND HEKETA ...... 215 TABLE 9.1 LIST OF ALL SPECIES IDENTIFIED ARCHAEOBOTANICALLY WITHIN THIS STUDY ...... 222 TABLE 9.2 IDENTIFIED FAMILIES AND SPECIES WITHIN ARCHAEOBOTANICAL REMAINS FROM TALASIU (TO- MU-2) ...... 241 TABLE 9.3 YIELD RATIOS FOR SPECIES IDENTIFIED AT TALASIU MODELLED USING COMPARATIVE SYSTEMS ...... 243 TABLE 9.4 LABOUR INPUTS FOR SPECIES IDENTIFIED AT TALASIU MODELLED USING COMPARATIVE SYSTEMS ...... 243 TABLE 9.5 STATISTICAL COMPARISON OF LABOUR INPUTS FOR GROUPINGS AT TALASIU IN TERMS OF MEAN DIFFERENCE MODELLED USING COMPARATIVE SYSTEMS ...... 243 TABLE 9.6 IDENTIFIED FAMILIES AND SPECIES WITHIN ARCHAEOBOTANICAL REMAINS FROM LEKA (J17) 245 TABLE 9.7 YIELD RATIOS FOR SPECIES IDENTIFIED AT LEKA MODELLED USING COMPARATIVE SYSTEMS . 246 TABLE 9.8 LABOUR INPUTS FOR SPECIES IDENTIFIED AT LEKA MODELLED USING COMPARATIVE SYSTEMS ...... 247 TABLE 9.9 STATISTICAL COMPARISON OF LABOUR INPUTS FOR GROUPINGS AT LEKA IN TERMS OF MEAN DIFFERENCE MODELLED USING COMPARATIVE SYSTEMS ...... 247 TABLE 9.10 IDENTIFIED FAMILIES AND SPECIES WITHIN ARCHAEOBOTANICAL REMAINS FROM HEKETA (TO- NT-2) ...... 248 TABLE 9.11 YIELD RATIOS FOR SPECIES IDENTIFIED AT HEKETA MODELLED USING COMPARATIVE EXAMPLE SYSTEMS...... 249 TABLE 9.12 LABOUR INPUTS FOR SPECIES IDENTIFIED AT HEKETA MODELLED USING COMPARATIVE EXAMPLE SYSTEMS ...... 249 TABLE 9.13 STATISTICAL COMPARISON OF LABOUR INPUTS FOR GROUPINGS AT HEKETA IN TERMS OF MEAN DIFFERENCE MODELLED USING COMPARATIVE SYSTEMS ...... 250

TABLE 9.14 COMPARISON OF MODELLED SYSTEM EFFICIENCY WITH RATIOS OF PRIMARY TO SUPPLEMENTARY SPECIES ...... 255

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List of Figures

FIGURE 5.1 FLOWCHART SHOWING METHODOLOGY FOR THE IMAGING AND RECORDING OF STARCH AND PARENCHYMA WITHIN THE REFERENCE COLLECTION...... 59 FIGURE 5.2 DIAGRAM SHOWING BASIC FEATURES OF STARCH GRANULE MORPHOLOGY ...... 65 FIGURE 5.3 BOX PLOT OF STARCH GRANULE LENGTHS WITHIN REFERENCE COLLECTION ...... 72 FIGURE 5.4 BOX PLOT OF STARCH GRANULE WIDTHS WITHIN REFERENCE COLLECTION ...... 72 FIGURE 5.5 BOX PLOT OF STARCH GRANULE HILUM POSITION TO LENGTH RATIOS WITHIN REFERENCE COLLECTION ...... 73 FIGURE 5.6 PLOT SHOWING DISCRIMINATION OF SPECIES WITHIN CENTRIC DATASET ACCORDING TO FIRST TWO CANONICAL VARIATES ...... 79 FIGURE 5.7 PLOT SHOWING DISCRIMINATION OF 15 SPECIES WITHIN ECCENTRIC DATASET ACCORDING TO FIRST TWO CANONICAL VARIATES ...... 79 FIGURE 5.8 CLASSIFICATION MATRIX FOR THE OVERALL CENTRIC DATASET, SHOWING HIGHEST

DISCRIMINATION OF COLOCASIA ESCULENTA, INOCARPUS FAGIFER, MORINDA CITRIFOLIA AND SPONDIAS

DULCIS (SPECIES LISTED VERTICALLY IN THE FIRST COLUMN ARE THE ORIGINAL SPECIES, AND THOSE LISTED HORIZONTALLY IN THE TOP ROW ARE THE SPECIES TO WHICH DFA CLASSIFIED GRANULES) .. 81 FIGURE 5.9 CLASSIFICATION MATRIX FOR THE OVERALL ECCENTRIC DATASET, SHOWING HIGHEST

DISCRIMINATION OF COLOCASIA ESCULENTA, CURCUMA LONGA, AND DIOSCOREA PENTAPHYLLA

(SPECIES LISTED VERTICALLY IN THE FIRST COLUMN ARE THE ORIGINAL SPECIES, AND THOSE LISTED HORIZONTALLY IN THE TOP ROW ARE THE SPECIES TO WHICH DFA CLASSIFIED GRANULES) ...... 81 FIGURE 5.10 BOX PLOT OF PARENCHYMA CELL LENGTHS OF TAXA IN THE REFERENCE COLLECTION ...... 87 FIGURE 5.11 BOX PLOT OF PARENCHYMA CELL WIDTHS OF TAXA IN THE REFERENCE COLLECTION ...... 88 FIGURE 5.12 PLOT SHOWING CLASSIFICATION OF PARENCHYMA WITHIN REFERENCE COLLECTION USING DFA ...... 89 FIGURE 5.13 DESCRIPTION OF VASCULAR BUNDLE ARRANGEMENTS WITHIN VEGETATIVE PARENCHYMA (FROM HATHER 2000) ...... 90 FIGURE 5.14 BOX PLOT SHOWING VASCULAR BUNDLE LENGTHS WITHIN REFERENCE COLLECTION ACCORDING TO TISSUE ARRANGEMENT ...... 94 FIGURE 5.15 FLOWCHART 1 USED AS AN IDENTIFICATION KEY TO IDENTIFY UNKNOWN PARENCHYMATOUS SAMPLES WHEN VASCULAR TISSUES ARE VISIBLE ...... 110 FIGURE 5.16 FLOWCHART 2 USED AS AN IDENTIFICATION KEY TO IDENTIFY UNKNOWN PARENCHYMATOUS SAMPLES WHEN NO VASCULAR TISSUES ARE VISIBLE...... 111 FIGURE 6.1 MAP SHOWING LOCATION OF ARCHAEOLOGICAL SITES INCLUDED IN THIS STUDY FROM TONGATAPU ...... 116 FIGURE 6.2 STRATIGRAPHIC DIAGRAM OF CULTURAL DEPOSITS WITHIN TALASIU TP2 ...... 120 FIGURE 6.3 STRATIGRAPHIC DIAGRAM OF CULTURAL DEPOSITS WITHIN LEKA TP2 ...... 122 FIGURE 6.4 STRATIGRAPHIC DIAGRAM OF CULTURAL DEPOSITS AT LEKA TP4 ...... 122 FIGURE 6.5 STRATIGRAPHIC DIAGRAM OF CULTURAL DEPOSITS WITHIN HEKETA TP3...... 124 FIGURE 6.6 CALIBRATION OF RADIOCARBON DATES FROM TALASIU (TO-MU-2), LEKA (J17) AND HEKETA (TO-NT-2) ...... 125 FIGURE 8.1 COMPOSITION OF OVERALL MACROBOTANICAL ASSEMBLAGE IN TERMS OF ABUNDANCE...... 139 xiii

FIGURE 8.2 BOX PLOT DEMONSTRATING MAXIMUM LENGTH COMPARISON OF ARCHAEOLOGICAL STARCH TYPE 1 WITH DIOSCOREA SPP...... 149 FIGURE 8.3 DISCRIMINANT ANALYSIS PLOT FOR CENTRIC DATASET SHOWING ELLIPSES (COLOURED DOTS REPRESENT REFERENCE SPECIES, BLACK DOTS REPRESENT ARCHAEOLOGICAL GRAINS) ...... 152 FIGURE 8.4 DISCRIMINANT ANALYSIS PLOT FOR ECCENTRIC DATASET SHOWING ELLIPSES (COLOURED DOTS REPRESENT REFERENCE SPECIES, BLACK DOTS REPRESENT ARCHAEOLOGICAL GRAINS) ...... 153 FIGURE 8.5 ARCHAEOLOGICAL AND REFERENCE STARCH: (A) ARCHAEOLOGICAL STARCH IDENTIFIED AS

ARTOCARPUS ALTILIS, (B) MODERN STARCH OF A.ALTILIS, (C) ARCHAEOLOGICAL STARCH IDENTIFIED

AS ALOCASIA MACRORRHIZA, (D) MODERN STARCH OF A.MACRORRHIZA, (E) ARCHAEOLOGICAL STARCH

IDENTIFIED AS AMORPHOPHALLUS PAEONIIFOLIUS, (F) MODERN STARCH OF A.PAEONIIFOLIUS, (G)

ARCHAEOLOGICAL STARCH IDENTIFIED AS BARRINGTONIA ASIATICA, (H) MODERN STARCH OF

B.ASIATICA, (I) ARCHAEOLOGICAL STARCH IDENTIFIED AS COLOCASIA ESCULENTA, (J) MODERN STARCH

OF C. ESCULENTA, (K) ARCHAEOLOGICAL STARCH IDENTIFIED AS CURCUMA LONGA, (L) MODERN

STARCH OF C.LONGA, (M) ARCHAEOLOGICAL STARCH IDENTIFIED AS CYRTOSPERMA MERKUSII, (N)

MODERN STARCH OF C.MERKUSII, (O) ARCHAEOLOGICAL STARCH IDENTIFIED AS DIOSCOREA ALATA, (P) MODERN STARCH OF D.ALATA...... 159 FIGURE 8.6 ARCHAEOLOGICAL AND REFERENCE STARCH CONT.: (Q) ARCHAEOLOGICAL STARCH IDENTIFIED

AS DIOSCOREA BULBIFERA (R) MODERN STARCH OF D. BULBIFERA, (S) ARCHAEOLOGICAL STARCH

IDENTIFIED AS DIOSCOREA ESCULENTA, (T) MODERN STARCH OF D.ESCULENTA, (U) ARCHAEOLOGICAL

STARCH IDENTIFIED AS DIOSCOREA NUMMULARIA, (V) MODERN STARCH OF D.NUMMULARIA, (W)

ARCHAEOLOGICAL STARCH IDENTIFIED AS INOCARPUS FAGIFER, (X) MODERN STARCH OF I. FAGIFER,

(Y) ARCHAEOLOGICAL STARCH IDENTIFIED AS IPOMOEA BATATAS, (Z) MODERN STARCH OF I.BATATAS,

(AA) ARCHAEOLOGICAL STARCH IDENTIFIED AS MUSA SP., (AB) MODERN STARCH OF MUSA SP., (AC)

ARCHAEOLOGICAL STARCH IDENTIFIED AS PIPER METHYSTICUM, (AD) MODERN STARCH OF

P.METHYSTICUM, (AE) ARCHAEOLOGICAL STARCH (CONTAMINANT) IDENTIFIED AS SOLANUM

TUBEROSUM, (AF) MODERN STARCH OF S.TUBEROSUM, (AG) ARCHAEOLOGICAL STARCH IDENTIFIED AS SPONDIAS DULCIS, (L) MODERN STARCH OF S. DULCIS...... 160 FIGURE 8.7 NUTRITIONAL COMPARISON OF SPECIES WITHIN THE GADIO ENGA PLANT PRODUCTION SYSTEM (DATA FROM DORNSTREICH 1974, 1978) ...... 164 FIGURE 8.8 NUTRITIONAL COMPARISON OF SPECIES WITHIN THE BELLONA ISLAND PLANT PRODUCTION

SYSTEM, SHOWING EXPONENTIAL TREND LINES FOR HORTICULTURAL AND SEMI-CULTIVATED TAXA (DATA FROM CHRISTIANSEN 1975) ...... 167 FIGURE 8.9 NUTRITIONAL COMPARISON OF SPECIES WITHIN THE ANUTAN PLANT PRODUCTION SYSTEM,

SHOWING EXPONENTIAL TREND LINES FOR PRIMARY AND SUPPLEMENTARY TAXA (DATA FROM YEN 1973B) ...... 171 FIGURE 8.10 NUTRITIONAL COMPARISON OF SPECIES WITHIN THE TONGAN PLANT PRODUCTION SYSTEM,

SHOWING EXPONENTIAL TREND LINES FOR HORTICULTURAL AND SEMI-CULTIVATED TAXA (DATA FROM MINISTRY OF AGRICULTURE AND FORESTRY 2001) ...... 174 FIGURE 8.11 NUTRITIONAL COMPARISON OF SPECIES WITHIN THE ONTONG JAVA PLANT PRODUCTION

SYSTEM, SHOWING EXPONENTIAL TREND LINES FOR PRIMARY AND SUPPLEMENTARY TAXA (DATA FROM BAYLISS-SMITH 1973, 1986) ...... 177

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FIGURE 8.12 LABOUR COMPARISON OF SPECIES WITHIN THE GADIO ENGA SYSTEM (DATA FROM DORNSTREICH 1974, 1977) ...... 180 FIGURE 8.13 LABOUR COMPARISON OF SPECIES WITHIN THE BELLONA ISLAND SYSTEM (DATA FROM CHRISTIANSEN 1975) ...... 181 FIGURE 8.14 LABOUR COMPARISON OF SPECIES WITHIN THE ANUTAN SYSTEM (DATA FROM YEN 1973B) 183 FIGURE 8.15 LABOUR COMPARISON OF SPECIES WITHIN THE TONGAN SYSTEM (DATA FROM MINISTRY OF AGRICULTURE AND FORESTRY 2001) ...... 185 FIGURE 8.16 LABOUR COMPARISON OF SPECIES WITHIN THE ONTONG JAVA PLANT PRODUCTION SYSTEM (DATA FROM BAYLISS-SMITH 1973, 1986) ...... 186 FIGURE 8.17 OUTPUT COMPARISON ACCORDING TO YIELD FOR SPECIES WITHIN GADIO ENGA SYSTEM (DATA FROM DORNSTREICH 1974, 1978) ...... 188 FIGURE 8.18 OUTPUT COMPARISON ACCORDING TO YIELD FOR SPECIES WITHIN BELLONA SYSTEM (DATA FROM CHRISTIANSEN 1975)...... 190 FIGURE 8.19 OUTPUT COMPARISON ACCORDING TO YIELD FOR SPECIES WITHIN THE ANUTAN SYSTEM (DATA FROM YEN 1973B) ...... 191 FIGURE 8.20 OUTPUT COMPARISON ACCORDING TO YIELD FOR SPECIES WITHIN THE TONGAN SYSTEM (DATA FROM MINISTRY OF AGRICULTURE AND FORESTRY 2001) ...... 193 FIGURE 8.21 COMPARISON ACCORDING TO YIELD FOR SPECIES WITHIN THE ONTONG JAVA PRODUCTION SYSTEM (DATA FROM BAYLISS-SMITH 1973, 1986) ...... 194 FIGURE 8.22 OUTPUT TO INPUT RATIO COMPARISON FOR ARCHAEOLOGICAL SPECIES WITHIN EACH SYSTEM IN TERMS OF CALORIES ...... 206 FIGURE 8.23 OUTPUT TO INPUT RATIO COMPARISON FOR ARCHAEOLOGICAL SPECIES WITHIN EACH SYSTEM IN TERMS OF PROTEIN ...... 207 FIGURE 8.24 OUTPUT TO INPUT RATIO COMPARISON FOR ARCHAEOLOGICAL SPECIES WITHIN EACH SYSTEM IN TERMS OF FATS (NOTE VERTICAL SCALE IS LOGARITHMIC) ...... 207 FIGURE 8.25 OUTPUT TO INPUT RATIO COMPARISON FOR ARCHAEOLOGICAL SPECIES WITHIN EACH SYSTEM IN TERMS OF CARBOHYDRATES ...... 208 FIGURE 8.26 COMPARISON OF AVERAGE NUTRITIONAL EFFICIENCY RATIOS FOR ALL SYSTEMS (NOTE VERTICAL SCALE IS LOGARITHMIC) ...... 208 FIGURE 8.27 NUTRITIONAL COMPARISON OF SPECIES IDENTIFIED AT TALASIU (TO-MU-2) ...... 210 FIGURE 8.28 NUTRITIONAL COMPARISON OF SPECIES IDENTIFIED AT TALASIU WITH EXPECTED ETHNOGRAPHIC SPECIES...... 211 FIGURE 8.29 NUTRITIONAL COMPARISON OF SPECIES IDENTIFIED AT LEKA (J17) ...... 212 FIGURE 8.30 NUTRITIONAL COMPARISON OF SPECIES IDENTIFIED AT LEKA WITH EXPECTED ETHNOGRAPHIC SPECIES ...... 213 FIGURE 8.31 NUTRITIONAL COMPARISON OF SPECIES IDENTIFIED AT HEKETA (TO-NT-2) ...... 214 FIGURE 8.32 NUTRITIONAL COMPARISON OF SPECIES IDENTIFIED AT HEKETA WITH EXPECTED ETHNOGRAPHIC SPECIES ...... 215 FIGURE 8.33 COMPARISON OF CALORIFIC EFFICIENCY OF ARCHAEOLOGICAL SPECIES FROM TALASIU...... 218 FIGURE 8.34 COMPARISON OF CALORIFIC EFFICIENCY OF ARCHAEOLOGICAL SPECIES FROM LEKA ...... 218 FIGURE 8.35 COMPARISON OF CALORIFIC EFFICIENCY OF ARCHAEOLOGICAL SPECIES FROM HEKETA ...... 219

FIGURE 8.36 MODELLED ARCHAEOLOGICAL SYSTEMS ACCORDING TO CALORIFIC EFFICIENCY VALUES FROM MODERN SYSTEMS ...... 220 FIGURE 9.1 TREND TOWARDS DECREASED SYSTEM NUTRITIONAL EFFICIENCY AND INCREASED SOCIAL COMPLEXITY AFTER LAPITA OCCUPATION AT TALASIU (2750-2650 CAL BP) ...... 259

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List of Abbreviations

BP years before present bsl below surface level bd below datum ºC degrees centigrade cal calibrated cm centimetres cm³ centimetres cubed DFA Discriminant Function Analysis hr hours ioa instance of activity km kilometres m metres m² metres square µm micrometre sg specific gravity

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Chapter 1 Introduction

Plants have played a critical role throughout history, enabling human migration, colonisation and the formation of complex societies. Many plant species have been biologically adapted to human exploitation through genetic manipulations that increase productivity and ease cultivation and transportation. The production and use of plants through horticultural practices was a critical stepping stone in the development of complex societies in human history. Plants enabled nutritionally diverse diets, but also provided useful materials such as cordage, fibre, thatch, cooking and storage vessels, and medicines to name a few. Plant production systems within the Pacific region are the result of transportation of plants and of concepts relating to them, as well as localised adaptation to specific island environments. Understanding the role that these systems played in stimulating and enabling episodes of human movement into and within Polynesia is critical for developing models of global migration patterns.

There are two prevailing views on human-environment interactions. The first of these argues that humans reacted passively to environmental change. This traditional perspective has been challenged by the view that humans act as agents of change, both reacting to and transforming the landscapes which they inhabit. Island environments represent microcosms of this ecosystem manipulation and adaptation, and the islands of the Pacific provide an important setting to test these views. Natural and cultural factors that affected the timing and speed of movement, settlement patterns, technological development and the transformation of island environments within the Pacific can arguably provide insights at a global scale.

This archaeobotanical thesis will focus on the links between plant utilisation and migration episodes within Western Polynesia as well as on the role of plants within the evolution of social hierarchy through agricultural development, which form core issues in debates about Pacific settlement processes. Specifically, this thesis will investigate the introduction and role of prehistoric crops in Tongan prehistory through a study of ancient plant remains found in Lapita and post-Lapita archaeological sites around Tongatapu.

Research aims and objectives A primary question examined in this thesis is whether early colonisers were dependent on introduced crops, or if human dispersal was fuelled predominantly by the exploitation of natural resources. On a global scale, human migration has traditionally been viewed as stimulated by the extraction of floral and faunal food resources within diverse environmental contexts. This stems from the causality concepts of ‘push’ and ‘pull’ factors at both place of origin and destination, which have been utilised by demographers, geographers and archaeologists alike for many decades (Anthony 1990; Lee 1966). The colonisation of diverse island landscapes within the Pacific Ocean, the largest of the Earth’s oceanic divisions at 165.25 million km², has been

1 recognised as an ideal setting to test this theory. The view that exploitation rather than adaptation drove episodes of Lapita migration into Remote Oceania was taken by Groube (1971). His ‘strandlooper’ concept for Lapita subsistence, later adopted by Best (1984), favoured marine exploitation over the utilisation of transported terrestrial crops and animals. This contrasts with the ‘transported landscape’ proposed by Kirch (1984) and others (Green 1991), in which a full suite of horticultural crops and techniques was brought by Lapita populations moving eastwards into Polynesia. These were then cultivated and intensified through adaptation of traditional practices within the varying high volcanic, limestone and atoll island settings of this region. The scale of ecosystem manipulation and the economic systems that fuelled this and later pulses of migration from the Western Polynesian homeland into Central and East Polynesia after 1500 BP are still contested. Global models such as the Ideal Free Distribution (IFD) from Human Behavioural Ecology (HBE) have been applied to account for modes of subsistence, new habitat suitability, and population density in predicting migratory behaviour (Kennett et al. 2006). Comparisons have also been drawn between the migration of food-producing people in areas such as the Pacific, Atlantic, Caribbean and the Mediterranean, suggesting that the insularity of island and coastal landscapes affects the rate and dynamics of colonising episodes (Dawson 2008; Keegan and Diamond 1987; Leppard 2014).

Disentangling local parameters and rates of change, as well as establishing how early subsistence evolved as locally-adapted cultures emerged is a crucial issue within Pacific archaeology that can be applied globally. Other archaeobotanical (Crowther 2005, 2009; Horrocks and Bedford 2004, 2010; Horrocks and Nunn 2007; Horrocks et al. 2009) and isotopic studies (Bentley et al. 2007; Field et al. 2009; Shaw et al. 2009; Valentin et al. 2010) have attempted to provide new proxy evidence on Lapita and post-Lapita subsistence, with research in Near and Remote Oceania. However, there has been very little direct evidence to corroborate the picture these data sketch. The analysis of both micro- and macrobotanical remains from archaeological sites has the potential to provide important new data to resolve the significance of human-plant production systems in the colonisation of the Pacific islands 3000 years ago.

Second, this study will examine whether archaeobotanical data can provide new evidence to examine the role of agriculture within the development of the maritime chiefdom in Tonga. Investigation of social complexity in Tonga has often assumed a causal link between the intensification of agriculture and the development of a complex chiefdom (Aswani and Graves 1998) or primary state (Clark et al. 2014) based on Tongatapu by 750 BP (Burley 1998). This approach stems from the foundational work of Childe (1925), Clarke (1952), Binford (1968) and Flannery (1965, 1968), among others, in Europe and the Americas. Simple environmental determinism and cultural systems theory have gradually evolved into ecological and evolutionary approaches that seek to explain the emergence of complex societies and the origins of agriculture through the dynamic that exists between people and their environment. These 2 approaches have generally moved away from unilinear trajectories for social complexity, and accept that multiple routes may be taken as a response to the same initial conditions. The ‘advantages’ of the adoption of agriculture did not necessarily ensure that this development would occur, and adaptive changes that might be viewed as regressive in the direction of less complex cultural forms also occurred (Binford 1968:331).

The islands of Polynesia present unique opportunities to study the links between tropical production systems, and agricultural intensification in non-industrialised societies. Testing of the ‘hydraulic hypothesis’ advanced by Wittfogel (1957), that draws a causal connection between the managerial requirements of complex irrigation and the development of complex socio-political structures, has been a dominant theme in Pacific research. Case studies from locations such as the Australs (Bollt 2012), Hawaii (Earle 1980, 1991, 2012; Ladefoged and Graves 2008; Lincoln and Ladefoged 2014; McCoy and Graves 2012; Sahlins 1958), the Marquesas (Addison 2006; Allen 2010; Earle 1993), and Futuna (Kirch 1982, 1994) have investigated the ties between investment in irrigation for wet taro production as landesque capital, and the development of political economies. The outcomes of these investigations challenged the idea of a direct link and demonstrated that territorial expansion was often the result of increasingly intensified dryland agricultural regimes when shorter-fallow and labor- intensive methods put pressure on the political elite to source other tracts of arable land. These more complex chains of causality question the uncritical imposition of generic hierarchical models of political economy that have often been derived from the largely discredited assumptions of Boserup (1965), Wittfogel, and the unilinear “Mesopotamian Model” of political economy, settlement pattern, and cultural evolution. ‘Expansion’, rather than ‘intensification’ of production to create food surplus therefore represents an alternative pathway to socio-political development both in the Pacific and elsewhere.

Research into dryland agricultural development on Tongatapu therefore presents an important case study that can be extrapolated and applied within global models linking production systems to social complexity. The dynamic relationship between known variables such as the environmental limitations for irrigation, the adoption of intensive dryland techniques, territorial expansion and centralised control of production through tributary systems in Tongan prehistory (Awani and Graves 1998; Burley and Connaughton 2007; Green 1973; Kirch 1988, 1994; Maude 1965; Spenneman 1986) are further explored in this thesis through a detailed chronology of plant introduction and cultivation. Specific questions are examined, such as whether the introduction of crops indicate trade and inter-island contact linked to socio- political development, and also if the creation of food surplus was likely enabled through intensification or expansion of production and the horticultural range. Did this process of agricultural development ultimately lead to changes in social hierarchy or were these variables inextricably linked? What factors were involved in these decision-making processes (i.e. 3 nutritional value, yield, labour investment)? This study sought to answer these research questions as well as to challenge existing assumptions which are primarily based on a lack of direct evidence for horticultural practices in prehistory, through systematic archaeobotanical analysis.

The two main objectives of this research were to construct a comprehensive reference collection for both starch and parenchyma from economic and supplementary plant species in Tonga, and to conduct a broad archaeobotanical survey of sediments from sites on the island of Tongatapu, from both Lapita-associated and post-Lapita contexts. Archaeobotanical techniques utilised here focus on the identification of both micro- and macrobotanical remains of plant storage organs. Plant microfossils such as starches, phytoliths and pollen have the potential to inform upon human interaction with their surrounding landscapes. In particular, these small remnants of plants can provide direct evidence for the role of plants within the diet and subsistence of a population. Starch is produced in the roots, tubers, fruits and seeds of plants, which are the main organs that are processed and eaten by humans (Torrence and Barton 2006). Additionally, macro-botanical remains such as charred, desiccated or water-logged vegetative storage parenchymatous tissue are also often diagnostic to species level (Hather 1991, 1994, 2000). These remains can enter into the archaeological record through either natural processes or through human intervention, such as intentionally growing or processing crops on site. An analysis of diagnostic morphological attributes of these remains was a crucial initial step towards enabling the taxonomic identification of material extracted from archaeological deposits, and built on the foundational work of others in the Pacific region (Babot 2003; Crowther 2001, 2005, 2009, 2012; Hather 2000; Horrocks and Barber 2005; Horrocks et al. 2004a, 2008, 2012, 2014; Loy 1994; Oliveira 2008, 2012; Paz 2005; Torrence et al. 2004; Wilson et al. 2010).

Three sites were targeted for this study: Talasiu (TO-Mu-2) dated to around 2750-2650 cal BP, Leka (J17 dated to 1300-1000 cal BP, and Heketa (TO-Nt-2) dated to 800-600 cal BP. According to Burley (1998), Tongan prehistory can be divided into four key periods: Lapita (2850-2650 BP), Plainware (2650-1550 BP), Formative Development (1550-750 BP) and Classic Tongan Chiefdom (750-150 BP). Archaeobotanical data from these three sites on Tongatapu has provided the basis for a revised chronology based on agricultural development rather than material culture, and changes the way that the relationship between subsistence and socio-political change is currently viewed in Tonga. It will be argued here that the early colonists of Tonga were primarily foragers, with minor elements of agriculture assisting subsistence. Agriculture was therefore not a later development in Tongan prehistory. Instead, reliance on the core cultigens likely grew as population size increased and new crops were introduced through trade and inter-island contact, ultimately allowing a state-level social hierarchy to emerge through the establishment and control of surplus resources. 4

Theoretical framework Basics of agroecology In order to address the two primary research questions within this study, a Human Ecological approach was taken to model and interpret the archaeobotanical data from Talasiu (TO-Mu-2), Leka (J17), and Heketa (TO-Nt-2). More specifically, these archaeological systems were analysed using techniques deriving from Agricultural Ecology or Agroecology, which aims to visualise cultivation as the creation of an ecosystem or agro-ecosystem (Tivy 1990). The management of crops and the environment through cultivation produces a habitat which allows the crop to realise its productive potential. Within this view, the agent becomes an essential ecological variable in the system, and the ‘ecosystem’ refers to social, cultural and economic contexts as well as the properties of the natural environment. It is argued that due to positive and negative feedback loops, whenever humans intercede they generate basic changes in the functioning of the system (Cox and Atkins 1979:57). Bayliss-Smith (1977) defines four types of productive efficiency that are calculated within this approach: indigenous efficiency (perceived output divided by primary input or cost in human effort to supply a population with goods to maintain and enrich it), exogenous efficiency (exported de-facto output divided by secondary input or cost to society as a whole of maintaining and enriching any enclave within it), technoenvironmental efficiency (Harris’ T) (total de-facto output divided by primary input), and total efficiency (total de-facto output divided by primary and secondary inputs).

Variables considered in these agro-ecosystems often relate to the mode of subsistence, labour inputs, yield, productivity, demography, as well as social and political constraints like surplus requirements. Previous applications of this approach in the Pacific have attempted to model production systems using ethnographic and historic data, and also island carrying capacity to refine population estimates (Bartruff et al. 2012; Bayliss-Smith 1977, 1978) and understand socio-political development (Lincoln and Ladefoged 2014). These studies vary in detail depending on the nature of data available, but often caution against strictly linking resource production to environmental restrictions or population pressure, as social factors also play a significant role in determining both population and modes of subsistence (Bayliss-Smith 1978). The degree of precision required to identify the ecological, economic and social constraints of agro-ecosystems ensures that archaeologists can only broadly hypothesise upon the interactions between natural and cultural variables. It is difficult to extrapolate whole prehistoric production systems from historic and ethnographic data, and how these changed over time.

Modern comparative systems and modelling archaeological systems Studies of modern agricultural production on Tongatapu have provided descriptions and data upon the nature of land ownership, cultivation techniques, cropping systems and fallow periods, and basic productivity within traditional subsistence subsidised by cash-cropping (Maude 1965; 5

Thaman 1976). This data is important, but cannot be assumed to accurately represent the nature of production throughout Tongan prehistory. The lack of diversity within modern production on Tongatapu required data collection from other Pacific dryland systems, to be able to model potential changes in yield and labour inputs resulting from varying cultivation techniques targeting different ranges of crops. Five systems from the Western Pacific were chosen, to provide a range of comparable data. They are the Gadio Enga of the New Guinea Highlands (Dornstreich 1977), Tongatapu (Ministry of Agriculture and Forestry 2001), Bellona Island (Christiansen 1975), Ontong Java (Bayliss-Smith 1973, 1977, 1986) and Anuta (Yen 1973b) in the Solomon Island Outliers. Data was collected on the range of plant species cultivated within each system, and the labour inputs and yields for each of these. These figures were used to calculate basic efficiency or rate of return ratios for each of these species in terms of nutritional and yield outputs to labour investment inputs, and to model archaeologically identified species within a range of different environmental, social and economic contexts. The geographic scale of each of these systems varied, but efficiency ratios are used to ensure comparability across all systems. Each system was characterised according to these variables (species diversity, nutritional diversity, labour diversity, and yield ratio diversity) rather than using loaded terminology such as ‘broad spectrum’ or ‘intensive’ that often does not capture the range of production techniques, decision-making and energy investment within different agro- ecosystems.

To model agricultural development in Tongan prehistory, it was deemed appropriate to keep the range of assumptions minimal to reduce the potential for error. Therefore an assessment of productivity was conducted through characterisation of archaeobotanical datasets using the ranges of yield and labour figures recorded from modern systems. Productivity is only one variable within an agroecological approach to production, but it can be placed within the social, ecological and economic context of each system to discuss how the range compared with expected outcomes based on previous research. It is clear that demographic, climatic and social factors would have impacted both the scale and intensity of agricultural production in the past, but these impacts can only be hypothesised within current research. The approach taken within this thesis therefore aimed to keep modelled assumptions to a minimum and redirect the focus of agricultural development away from the scale of production, towards a discussion of decision-making based on system nutritional efficiency and founded on data highlighting the timing of crop introductions and their use.

Thesis organisation This study of agricultural development in Tongan prehistory is broken down into two main components related to the objectives of this research. After a brief review of Tongan agriculture within the Pacific context (Chapter 2), the first component will focus on the development of a comparative collection of micro- and macrobotanical remains for Tonga. Chapters 3 and 4 6 assess the feasibility of applying the analysis of starch and vegetative parenchyma to questions regarding diet and subsistence. The biology, morphology, taphonomy and contamination potential of each of these remains was reviewed in order to gauge both the preservation and identification potential of archaeobotanical material. Chapter 5 outlines the methodology to create a reference collection for this study including species selection, field collection, and laboratory processing. Sample preparation for imaging using light microscopy and Scanning Electron Microscopy (SEM) is described, and the collection of morphometric data through the use of image analysis software. Finally, tools such as replicable multivariate statistical classifications and identification keys are provided that characterise the morphology of starch and vegetative parenchyma and will enable the identification of unknown samples from archaeological deposits. The development of this comparative collection will enable the key research questions concerning the role of plants within the colonising process and the development of social hierarchy in the Tongan archipelago to be addressed.

The second component of this thesis deals with the application of micro- and macrobotanical techniques within archaeobotanical research on Tongatapu to provide a chronology for crop introductions and agricultural development. Chapter 6 introduces the three sites selected for analysis, Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2), and details the methods employed to excavate and extract archaeobotanical remains from cultural deposits. The establishment of laboratory protocols for the extraction and analysis of both starch and macrobotanical remains such as charred endocarp and parenchyma is explored in Chapter 7. This borrows from and also builds on the range of protocols already used in archaeobotanical studies in the Pacific region. The results of extraction and identification processes from the three sites are outlined in Chapter 8, along with the cost-benefit analysis of five comparative plant production systems from the Western Pacific and the archaeological production systems from each site. Chapter 9 discusses the characterisation and modelling of data from each of the archaeological sites included in this study. The feasibility of modelling past production systems using a Human Ecological and Agroecological approach is discussed, along with a comparison of the expected and modelled outcomes of analysis and how modelled production can be tied to social complexity. Chapter 9 also discusses the timing of individual crop introductions into Tonga using the data provided within this study and a new chronology for agricultural development in Tongan prehistory is outlined. Chapter 10 concludes this thesis with a summary of the outputs of this research: the development of a comprehensive comparative collection for Tonga that has applications for the wider Pacific, and an alternative perspective on agricultural development in Tonga based on proxy and direct evidence for crop cultivation and use. Recommendations for future research that will build on these outputs through further application of archaeobotanical techniques such as starch and charred parenchyma analysis within research in Tonga and Pacific region is also highlighted and emphasis is placed on the

7 potential contribution these remains can make towards answering questions about diet, subsistence and social complexity in prehistory.

Chapter 2 Tongan Agriculture in the Pacific Context Research to date Research into Tongan prehistory has thus far been oriented towards the study of artefacts and faunal remains. Although a range of archaeological and ethno-historic research has already been carried out in Tonga, none of these studies have so far addressed the role of agricultural change within cultural development in the island group. Early research was directed towards gathering information about the first colonisers, known as the Lapita culture, who had made their way from Southeast Asia into Melanesia and finally into Western Polynesia by 2800 BP (Burley 1998:349; Burley and Connaughton 2007; Burley et al. 2001). Very little is known about the role of horticulture within Lapita subsistence, especially in Western Polynesia. However, there are numerous studies of the wild food components of colonising diets such as shellfish, birds, reptiles and fish (Burley 1998; Burley and Connaughton 2007; Burley and Dickinson 2001; Groube 1971; Kirch and Dye 1979; Poulsen 1987; Spenneman 1986, 1989; Steadman, Pregill and Burley 2002). Archaeologists have been divided over whether a ‘strandlooper’ economy was employed that focussed on the collection of these natural coastal resources (Best 1984; Groube 1971); or a subsistence dominated by well-developed agricultural practices (Green 1979; Kirch 1997). The interpretation of the archaeological record is biased towards marine- based foraging due to the dominance of midden remains within Lapita cultural deposits (Davidson 1979:93; Davidson and Leach 2001), and current evidence suggests that agricultural activities were initially of only secondary importance (Burley 1998:355; Poulsen 1987:253-5; Spenneman 1989).

The signature dentate-stamped pottery has become the cultural marker for the Lapita, but after 2650 BP a trend of undecorated pottery (Plainware) can be seen in the archaeological record (Burley 1998:359). This is viewed as part of the development of Western Polynesian culture within Tonga, and was part of a suite of changes that also included expansion further inland and into off-shore islands. Kirch (1984) argued that expansion was an indication that population density also began to increase dramatically. Kirch calculated that roughly all of the arable land on Tongatapu would have come under agrarian use between one to two millennia after colonisation (1984:222). These, and earlier calculations were based on agricultural practices involving shifting cultivation with relatively short fallow periods, as recorded by ethnographers after European contact (Beaglehole and Beaglehole 1941; Green 1973). During this same period, after 2650 BP reliance on lagoon-based marine resources such as shellfish also decreased markedly (Spenneman 1989).

Between 1500 BP and around 750 BP, very little is known about Tonga’s past. Oral traditions and genealogy suggest that the first Tu’i Tonga was appointed at around 1000 BP, although evidence of a widespread and integrated maritime chiefdom within the archaeological record is difficult to confirm prior to 500 BP (Burley 1998:375). Janet Davidson coined the term ‘the Dark Age’ for this gap, citing the lack of archaeological sites until late in the first millennium AD when monumental architecture associated with the development of a complex chiefdom emerges (Davidson 1979:94-5). More is known about the Tongan maritime chiefdom after 750 BP and the control of land by this central polity based on Tongatapu (Burley 1998; Clark et al. 2008). A complex social hierarchy existed at European contact, culminating in three paramount chiefly titles known as the Tu’i Tonga, the Tu’i Ha’a Takalaua, and the Tu’i Kanokupolu lines.

To date, there is currently no direct archaeological evidence for agriculture in Tongan prehistory. Historical linguistics indicates a Proto-Oceanic lexicon containing the basic suite of Oceanic cultigens (Kirch 1997:206-207), which is assumed to have been transported throughout Western Polynesia by the Eastern Lapita cultural complex. Thus it is inferred that the Lapita colonisers had an economy in which agriculture played some continued role. In the late 18th century the Tongan language had names for a wide range of root and tree crops, although the core cultigens grown were yam (Dioscorea alata), giant taro (Alocasia macrorrhiza), taro (Colocasia esculenta), sweet potato (Ipomoea batatas), coconut (Cocos nucifera), plantain and bananas (Musa spp.), and breadfruit (Artocarpus altilis) (Burley and Connaughton 2007:182). Ethno-archaeological observations have been made of multi-cropping using intensive dryland field systems that are suited to the high limestone islands and lack of streams for irrigation (Kirch 1984). Agricultural features related to these systems, such as stone structures, can also still be seen in some locations in the Tongan landscape. Although field systems have been mapped on the outer islands, such as Niuatoputapu (Kirch 1988), no radiocarbon dates are associated with these features. Based on this information, it has been assumed that a changing environment, population increase and subsequently reduced food returns forced post-Lapita Tongan populations to rely more heavily on horticulturally produced food, eventually manifesting in an increasingly hierarchical society (Spenneman 1986:3).

Geographic and climatic limitations to agricultural modelling The Kingdom of Tonga comprises more than 170 islands within the South Pacific Ocean, and is located from 15°30’ to 22° 20’ S latitude and between 173°00’ and 177° 15’ W longitude. The islands formed on the crest of the Tongan Ridge, bordering the Tonga Trench at the Pacific Plate Boundary (Roy 1990). Tongatapu is within the southern block of this ridge and is made up of Pliocene and Pleistocene limestone that reaches a maximum height of 65-70m above sea level in the southeast, and is little above sea level in the northwest due to a tilt in topography. As a result, the southern windward shores are composed of near-vertical cliffs and represent the most rugged topography on an island that is largely flat to gently undulating (Roy 1990; Maude 10

1965; Thaman 1976). There are no permanent streams on the island; free-draining porous soils above the limestone geological base which enable water to travel to the sea through underground channels. The location of the island of Tongatapu near the Tropic of Capricorn results in a mean annual temperature of around 23°C, humidity of up to 79%, and mean annual rainfall of 187cm (Thaman 1976). Seasonal variation in temperatures is low, only fluctuating by 4°C, but rainfall varies from around 8.4cm in the driest months to 24cm in wet months; however, there can be considerable variability in rainfall from year to year. Whilst there are some seasonal and annual fluctuations in climatic conditions, these do not vary significantly across Tongatapu due to the absence of significant topographic differences.

The geography and climatic conditions on Tongatapu both aid and hinder the modelling of past production systems. Unlike the Hawaiian archipelago, where dryland agricultural systems are often constructed and managed within distinct ecological zones (Lincoln and Ladefoged 2014), the raised limestone island of Tongatapu has little variety within ecological niches and so has no distinguishable boundaries for the utilisation of diverse production techniques (Maude 1965). One of the only observed variables affecting the geographic distribution of crop production is soil type. Two main soil types are present within the archipelago, the kelefatu soils which are very friable and fertile volcanic soils and vary in texture from loamy sand to clay, and the tou’one sandy soil that is present at low elevations close to the sea (Maude 1965; Thaman 1976). Tongatapu has both of these, but the kelefatu is the most prevalent- covering 90% of the landmass. Yams (Dioscorea spp.) do not grow well within small areas of tou’one soils; the sandy fast-draining soils are instead better suited to the cultivation of sweet potato (Ipomoea batatas). Gibbs (1967) distinguished two sub-categories between the upland or kelefatu soils. The first of these, ‘Lapaha clays’ are predominant in Eastern Tongatapu and around the capital Nuku’alofa. The other, ‘Vaini clays’ cover most of the uplands in the west. The two soils are very similar in texture and composition yet cultivators in modern times understand the differences in fertility resulting from use and manipulation of these soils, and the limitations of particular cropping and fallow practices within these (Maude 1965). It is therefore possible that almost all of the available arable land on Tongatapu, around 224.14km² or 55398 acres, could be cultivated using much the same shifting dryland agricultural techniques, at any point in time. Early censuses estimate that around 7, 308 people were living on Tongatapu by 1891(Burley 2007), while estimates for pre-contact populations for the whole archipelago vary from 29700 (Maude 1965) to 40000 (Kirch 1984, 1988).

These two sets of data (geographic variation and historic population figures) have been used to model carrying capacity during Tongan prehistory by Green (1973; Bayliss-Smith 1978), where populations were supported by per capita acreages of 1.6-2.0 depending on high to low intensity production. The standard populations were calculated for these intensity levels and varying proportions of arable land under cultivation, and suggest that under the most intensive 11 system (1.6 acres per person) the entire arable land could have supported 346200 people (Green 1973:70). Based on these figures, and ethnographic descriptions of cultivation systems within the landscape on Tongatapu, Green argues that the best population estimate for the 18th century would have been 15000-17000, under conditions of 1.8-2.0 acres per capita in a cultivation cycle of 8-10 years through bush fallowing agriculture (1973:72). Furthermore, this figure could easily have been attained up to 1000 years before, but this does not indicate that the population reached this peak and then stayed at this level. Instead, Green (1973:73) suggests that following this peak the pressure on land and resources triggered reduction mechanisms such as warfare to restrain further growth. Green also argues that reduction mechanisms do not necessarily begin after carrying capacity has been reached, but instead when figures of closer to 60-80% capacity are attained. These figures set an upper limit on population on Tongatapu of between 18,000 to 24,000 people and suggest population may have been higher in the past than at contact.

More recently, Burley (2007) modelled population estimates for the three key phases in Tongan prehistory (Lapita, Plainware and Classic Chiefdom). Previous estimates (Green 1973; Kirch 1984; Maude 1965; Walsh 1970), settlement patterns and analysis of material culture (Burley 1999; Spenneman 1987) were utilised to predict the likely rate of population increase over time. Given a founding population of around 100 people, a conservative population growth rate of 0.003, and a need for approximately 2 acres of productive land per person, the population at the end of the Lapita period within the archipelago could have been as high as 600-700 people (Burley 2007). Later during the Plainware and Ancestral Polynesian phases, around half the projected maximum land could have been under production with agricultural field systems by AD 400, which may have stimulated long-distance voyaging and exchange networks further east. Finally late prehistoric populations were predicted to have peaked around 18, 467 on Tongatapu alone (Burley 2007). The difference between past estimates and these recent figures indicate that there are varying opinions upon the potential agricultural productivity of land on Tongatapu, rates of population growth, as well as the acreage needed to support increasing populations.

Ethno-historic accounts of plant cultivation in Tonga: Early explorers and missionaries have provided vital details about Tongan subsistence practices at European contact. Most relevant to this study are descriptions of land tenure, and the cultivation of crops within the plantations they encountered throughout the archipelago. In addition, these sources mention the most important economic species including cultigens and wild species that were made use of in times of famine or hardship. The list of crops witnessed by Le Maire in 1616 (1967), Abel Tasman in 1643 (1776), Cook in 1773 and 1777 (in Beaglehole 1969; Cook 1785), La Perouse in 1788 (1799), Labillardiere in 1793 (1800), Wilson (1799), Mariner in 1791 (Martin 1991), Waldegrave in 1830 (1833), Hemsley (1894), Gifford (1929), Beaglehole and Beaglehole (1941), and others on Tongatapu, is vital to the development 12 of a comprehensive understanding of late prehistoric and contact period agriculture in Tonga. It is also possible with such information to assess the temporal changes in reliance on certain crops. Other researchers have established the value of using ethnographic data as a guide to understanding the past (David and Kramer 2001, Wylie 1985). Ethnohistoric information is used in this thesis to create baseline data for the selection and development of a comparative collection in this study, as well as generating a picture of the environmental and social limits of production in Tonga under the Tu’i Tonga chiefdom in the historic period.

Plantations Information can be gathered from these sources on the nature and organisation of agricultural plantations, within which the majority of cultivated crops were grown. Importantly, early observers also commented on the division of land and production according to status differentiation. The first European explorer to reach Tongatapu was Dutch explorer Abel Tasman in 1643 (1776), followed by Cook in 1773 and again in 1777 (in Beaglehole 1969; Cook 1785). Tasman (1776), having observed the bounty of Tongatapu through gifts and trade of pigs, poultry, coconuts, plantains, bananas, yams and other roots, went onshore and noted the layout of plantations in neat squares within which these crops were cultivated. During his second voyage in 1773, Cook (in Beaglehole 1969; Cook 1785) explored the islands within the Vava’u, Ha’apai and Tongatapu groups, and described his encounters with the islanders and excursions onshore. Cook, his officers, and onboard naturalists commented extensively on the distribution of plantations across the various islands, and the nature of the crops being cultivated (Cook in Beaglehole 1969; Cook 1785). On the main island of Tongatapu, Cook (1785) observed the layout and functional divisions within the individual plantations. The botanical knowledge of a naturalist on d Entrecasteaux’s voyage to the Vava’u Group, Jacques Labillardiere (1800), enabled the accurate identification and description of the various crops within these plantations, along with their domestic or economic purposes. On the larger island of Tongatapu, multi-cropping seemed to be the common method of cultivation within the individual plantations, whereby a variety of food plants are grown together within plots.

A surgeon and naturalist on Cook’s second and third voyages, Mr Anderson, travelled more extensively inland from the coast of Tongatapu and made further notes upon the distribution of cultivated areas. Eastward of their base near the Fanga ‘Uta lagoon, there was a lack of uncultivated land for nearly two miles (Anderson in Cook 1785:288). However, further west most of the country was composed of fenced plantations (1785:314). Johann Reinhold Forster was another naturalist on Cook’s second voyage whose journals, alongside those of his son Georg, described the cultivation of fruits and vegetables within organised plantations (Forster 1777, 1778). Georg noted that the coral rock, thin soils and lack of groundwater ensured that those inhabiting Tongatapu were, in his opinion, forced to labour more greatly than the Tahitians to produce food (Forster 1777). He argued that this accounted for the regularity of the 13 plantation distribution, and the accurate division of land (Forster 1777). Johann (Forster 1778:223) also observed that the whole of Tongatapu was highly cultivated and seemed to be private property, the boundaries of which were fenced. Tree crops also commonly bordered or divided plantations (La Perouse 1799:171).

Cook (in Beaglehole 1969) noted that there were differences between the produce of the plantations reserved for the chiefly elite or ‘first rank’, and the commoners. Similarly, Captain Wilson (1799) commented on the links between land tenure arrangements and the layout of individual plantations. A map was produced from the circumnavigation of Tongatapu, highlighting the occupied and cultivated areas of the island, and the traditional names for these locations. Wilson also noted the use of the term ‘abey’ [abi] for these plantations or plots of land (Wilson 1799:101), which is still in use today but under different tenure conditions.

Land tenure Complex social hierarchy controlled the division and use of land for agriculture on Tongatapu in the 18th and 19th Centuries. Accounts from this period describe how this status differentiation resulted in different conditions for land ownership. These details are essential for reconstructing the influence that the state-level social hierarchy may have had on crop introductions and associated horticultural production in the past. Captain Waldegrave (1833) was informed during his visit in 1830 that the island was divided into 13 portions, with a chief being the proprietor of each. He was told that chiefs could, and often did, displace residents on the land, and these chiefs retained a claim to a portion of the agricultural produce (1833:185). This portion was argued by Waldegrave to be claimed in the absence of an official ‘taxation system’. In addition, the kings and higher chiefs reserved a portion of the land itself for their own agricultural production. E. W Gifford spent around nine months in Tonga as a member of the Bayard Dominick Expedition of Bernice P. Bishop Museum in 1920–21. Gifford noted that in the past all land and its products were regarded as the property of the Tu’i Tonga (Gifford 1929:102). Within these lands, those within the domain of the Tu’i Kanokupolu were regarded as his property, but also subject to the demand of the Tu’i Tonga. Likewise, a similar relationship was continued between the lesser chiefs and the Tui Kanokupolu. Both Cook (1785) and Mariner (in Martin 1991) noted similar arrangements. Gifford also argued that there seems to have been no communal lands in Tongan prehistory, but at the time of his visit there were some clear examples of modern communal tenancy (1929:176). On Lifuka Island, a particular tract of land or api belonged to the Queen, but in 1920 it was used as a communal field by the inhabitants of the village of Pangai, each household having a small section to plant sweet potatoes (1929:176). During harvesting, a portion was then given in return to the Queen.

All vegetable and animal products within the territory directly controlled by the Tu’i Kanokupolu were tapu, or off-limits, for anyone but that high chief (Gifford 1929:104). Two

14 petty chiefs were appointed to continuously check that the common people were not utilizing those products without permission. The titles of these positions were the Tu’i Pangai and the Tu’i Sinoieiki. During their rounds of the territory, these officers would mark fine bunches of cooking bananas, breadfruit, yams, sweet potatoes that were suitable for the Tu’i Kanokupolu, by putting a sharp stick into the base of the tree or hanging coconut leaves from the branches (1929:104; Mariner in Martin 1991). In the case of yams or sweet potatoes, a stick was placed into the ground near the crops. These officers also reported to the chief and through him to the Tui Kanokupolu, upon the amount of crops planted by the farmers, and whether or not they were sufficient. However, this only applied to the territory of the Haa Ngata Motua lineage. A range of terms for other roles within Tongan society were also recorded by Gifford. For example, ‘pule fonua’ was the name for the rulers of the land, or chiefs who control food supply (1929:104).

Festivals Related to these intricate systems of land ownership was a yearly pattern of feasting and tributes based on the produce of plantations. According to early accounts, these were important for the redistribution of food surpluses and reinforced the ownership and land and production by elites within Tongan society. The first fruits of the yearly yam harvest or the first of any catch of fish or other food had to be presented to the chief and to the Tui Tonga before it could be partaken of by the producers (Gifford 1929:103). Tributes were carried from as far afield as Vava’u. A select group of people were appointed as petty officers in charge of supplying these food articles to the Tui Tonga, behind which Gifford believes is a form of religious sanction (1929:103). These tributes were “virtually made to the gods, but were made to them through the Tui Tonga who was treated like a god” (1929:103). William Mariner was a resident upon the island of Tongatapu for several years from 1806 to 1810, and recounted his experiences to John Martin which were originally published in 1817. During this time, he observed the cultivation of breadfruit, coconuts and yams on a daily basis, and so could provide detailed descriptions of horticultural practices throughout the seasons. He witnessed the ‘First fruits’ or ‘inasi ceremony, and perceived these offerings of the early yam harvest to be a means of insuring the “…protection of the gods and productions of the earth, of which yams are the most important.” (Mariner in Martin 1991:381)

In preparation for the ceremony, a particular variety of yams were planted about a month before the regular crop (Mariner in Martin 1991:381), most likely those described today as the ‘Early yam’. As soon as this crop arrived at a certain state of maturity, a message was sent to the Tui Tonga that the yams are fit to be harvested. The Tui Tonga then appointed a day for the ceremony— usually around 10 days later. The day before the ceremony the yams were dug up and wrapped in Pandanus leaves, and other provisions such as fish, kava root and mahoa (arrowroot) are prepared (Mariner in Martin 1991:382). On the day of the ceremony the harvest 15 was brought to the malai or meeting place of the chief of the plantation, and then the procession carried onto the grave of the last Tui Tonga to receive a ‘blessing’, before returning to the malai. The yam harvest was then divided into portions. Half was allocated to the king, one quarter was dedicated to the gods and so appropriated by the priests, and the remainder was given to the Tui Tonga (Mariner in Martin 1991:382). Mariner thus commented on the generosity of the Tongan people, particularly those of the lower ranks, whereby it was “…so much the custom of Tonga to make liberal and profuse presents that the people generally either feast or starve.” (Mariner in Martin 1991:385)

In addition to the ‘inasi festival, several other ceremonies were carried out during the harvests to ensure productive success. Mariner describes the ‘tow-tow’ [tau tau] ceremony whereby offerings of yams, coconuts and other vegetable produce was made to A’lo A’lo, the god of weather, in particular but also to other gods (Mariner in Martin 1991:385). These concessions were made for the purpose of ensuring a continuation of favourable weather and soil fertility. The ceremony was performed initially when the yams were approaching maturity in early November, and then repeated every ten days seven or eight times. The produce was piled into three mounds, one of which was given to the gods and the remaining two were given to the chiefs and their households. After the ceremony, the pile dedicated to the gods was then divided between the attendees of the festivities. Another form of feasting was the ‘pongipongi’ which involved the presentation of food and kava to the Tu’i Kanokupolu by chiefs of the Haa Ngata Motua and Haa Hatea lineages several times a year (Mariner in Martin 1991:99). In general, the Tu’i Kanokupolu was regarded as the ‘working king’ who oversaw the planting and other activities for the high king or Tu’i Tonga (Gifford 1929:99). The first Tu’i Kanokupolu Ngata was sent to Hihifo by the Tu’i Haa Takalaua to supervise agriculture and fishing, passing the produce to the Tu’i Haa Takalaua, then to the Tu’i Tonga, and the tradition continued until the independent powerbase in Hihifo eventually overthrew the Tu’i Tonga.

Cropping cycle Importantly for this study, a small number of European explorers and scientists also commented on the seasonality and production techniques used to cultivate and harvest crops produced within these plantations on Tongatapu. These shed light on the restrictions upon production of annual crops such as taro (Colocasia esculenta) over perennial tree crops such as breadfruit (Artocarpus spp.) or bananas and plantains (Musa spp.). Anderson (in Cook 1785) argued that the tropical climate of Tonga would have resulted in a fast turn-over of crops within the plantations. He commented:

The quick succession of vegetables has been already mentioned; but I am not certain that the changes of weather, by which it is brought about, are considerable enough to make them perceptible to the natives as to their method of life, or rather that they should be

sensible of the different seasons. This perhaps may be inferred from the state of their vegetable productions, which are never so much affected, with respect to the foliage, as to shed that all at once; for every leaf is succeeded by another, as fast as it falls, which causes that appearance of universal and continual spring round here. (Anderson in Cook 1785:330).

Likewise, Cook himself described the difference in production between his visits. It was noted that the stocks were replenished despite arriving back again sooner than expected, but the breadfruits which had been the only original produce available for purchase were replaced with yams and plantains (1785:272). Large areas that had been fallow previously were transformed into plantain fields. These observations demonstrate the relatively quick succession of the seasons, in terms of the vegetables produced on Tongatapu at different times of the year (in Beaglehole 1969). Seasonal fluctuations were also commented on by La Perouse (1799:171), who thought that the low islands in this group likely experienced drought at some stage during the year and commented on the necessity of watering fields. In contrast, on the island of Uoleva further north in the Ha’apai group, water sources enabled irrigation.

Summary From these various accounts, it is clear that some crops were used consistently from first European contact in the mid 18th century, right through until the early 20th century. The staple cultigens included Elephant ear taro (Alocasia macrorrhiza), breadfruit (Artocarpus altilis), coconut (Cocos nucifera), taro (Colocasia esculenta), winged yams (Dioscorea alata), the lesser yam (Dioscorea esculenta), sweet potato (Ipomoea batatas), plantain and bananas (Musa spp.), and Polynesian arrowroot (Tacca leontopetaloides). Other fruits, nuts and tubers were also cultivated to some degree and supplemented these crops. These included the shaddock (Citrus maxima), turmeric (Curcuma longa), melons (Cucumis melo), Tahitian chestnut (Inocarpus fagifer), Indian mulberry (Morinda citrifolia), pandanus (Pandanus tectorius), kava (Piper methysticum), sugarcane (Saccharum officinarum), Otaheite apple (Spondias dulcis), Malay apple (Syzygium malaccense), and the tropical almond (Terminalia catappa). Finally, a small range of naturalised plants were utilised for their edible fruits and tubers such as bitter yam (Dioscorea bulbifera), the giant swamp taro (Cyrtosperma merkusii), and the stink lily (Amorphophallus campanulatus). The majority of these crops in the proto-historic era were grown in plantations that surrounded small villages. Each household usually had the use of an individual plantation for their own subsistence needs, however the land itself could be owned by the chief or higher chiefs such as the Tui Kanokupolu or Tui Tonga (Gifford 1929:102, Waldegrave 1833:185). The land belonging to these chiefs was monitored by several officers appointed by the person holding one of these titles, and crops could be allocated to them upon maturation and harvesting (Gifford 1929:104). Related to these systems of land ownership was a

17 yearly pattern of feasting and tributes that were brought from as far afield as Vava’u (Mariner in Martin 1991:385).

The internal organisation of plantations varied, depending on the individual tastes and choices of the households utilising them. However, most households relied heavily on yams, multi- cropping them with sweet potato and taro in mounds in the centre of plantations (Beaglehole and Beaglehole 1941:43-44, Gifford 1929, La Billardiere 1800:366). Arboricultural crops such as plantains, bananas, and breadfruit were grown bordering the outer limits of the plantations (La Billardiere 1800:378, Wilson 1799:240). These trees were often left to mature and harvested each season as the fruits ripened, while the tuberous plants were replanted until the soil in the plot could no longer sustain growth and the plot was abandoned for a time (Cook in Beaglehole 1969; Cook 1785:271-2). After the introduction of new crops by European settlers such as manioc and corn, agricultural practices were forced to change to suit the requirements of these cultigens and there was a reduced reliance on traditional cultigens (Beaglehole and Beaglehole 1941). Despite this shift, these crops recorded by European explorers in the 18th and 19th Centuries mostly continue to be grown today.

These ethnohistoric details provide essential background information to meet the two main objectives of this thesis. The lists of species recorded within early accounts provided baseline data for the construction of a comprehensive reference collection of economic and supplementary plants used in Tongan prehistory. Likewise, the descriptions of production techniques, plantation layouts, feasting and tribute systems form a picture of agricultural production in the proto-historic period at the height of the state prior to European influence. These details enable some insight into the efficiency of this system in both productive and social terms, and against which past systems can be compared to link production to social complexity.

Tasman Cook Forster Perouse Labillardiere Wilson Mariner Orlebar Waldegrave Gifford Beaglehole Species 1643 1773-77 1773 1780s 1793 1797 1817 1830 1833 1929 1938 Alocasia macrorrhiza X X X Amorphophallus paeoniifolius X Artocarpus altilis X X X X X X X X X X Benincasa hispida X Broussonetia papyrifera X X X X X Citrus maxima X X X X Cocos nucifera X X X X X X X X X X X Colocasia esculenta X X X X X X X Cordyline fruticosa/terminalis X Curcuma longa X X Cucumis melo X Dioscora alata X X X X X X X X X X Dioscorea bulbifera ? X ? Dioscorea esculenta X ? X ? X Ficus tinctoria X Hibiscus manihot X Inocarpus fagifer X X X Ipomoea batatas X X X X Morinda citrifolia X Musa sp. (bananas) X X X X X X X X X X X Musa sp. (plantains) X X X X X X X Pandanus tectorius X X X Piper methysticum X X X X X X X X Pritchardia pacifica X Saccharum officinarum X X X X X X Spondias dulcis X X X Syzygium malaccense X Tacca leontopetaloides X X X

Table 2.1 List of species recorded in early ethno-historic accounts from Tonga. 19

Archaeobotany of cultigens in the Pacific The discussion of agricultural change in Tongan prehistory is the product of the direction of past research, rather than a paucity of evidence for this floral element of subsistence within the Tongan archaeological record. Microfossil research in other locations has demonstrated the applicability of this analysis towards identifying the presence of crops and the nature of their use within archaeological contexts (Fullagar et al. 1998, 2006; Horrocks et al. 2004; Horrocks, Grant-Mackie and Matissoo-Smith 2008; Loy et al. 1992; Piperno 2009; Summerhayes at al. 2010, among others). Similarly macrobotanical analysis has been used to provide direct evidence of plant cultivation and domestication in the New and Old Worlds, and the Indo- Pacific (Fuller 2007; Oliveira 2008; Paz 2001; Thompson 1994; Ugent et al. 1981).

Unlike archaeological investigations in the Old World, archaeobotanical techniques have not been consistently applied in the Pacific region. This is due to several factors. The first of these is that the preservation of plant remains in tropical climates is more variable than in the cooler or arid northern latitudes (Paz 2005). Second, the agricultural practices of most prehistoric Pacific communities revolved around the production and consumption of starchy tree nuts, roots and tubers, rather than small seeds such as wheat or rice which have been the traditional focus of archaeobotany (Hather 1994:51). Finally, many of the main domestic plants such as taro (Colocasia esculenta), bananas (Musa spp.), and breadfruit (Artocarpus spp.) are not genetically or phenotypically identifiable from wild populations (Fairburn 2005a). Consequently, archaeobotanical investigations in the tropics have often been based on chance finds (Paz 2005; Glover 1979). Recently, rock shelters, caves, and waterlogged sites have provided new site types for the preservation of plant remains in tropical climates (Glover 1979; Oliveira 2008; Paz 2001, 2005; Fairburn 2005a). Analysis of those remains that have been recovered have suggested that many tropical plant production systems are a poor fit within the traditional typologies of foraging or farming using wild or domestic species, and so the whole global system of agricultural classification has subsequently come into question (Fairburn 2005a).

These technical and theoretical issues have complicated the interpretation of plant use within the archaeobotanical record in the Pacific. Here, the goal is to review the antiquity of plant use within the Pacific economy, through focusing on the identification of macro- and microbotanical remains of edible root, tuber, and tree crops. Clearly domestication cannot be sufficiently discriminated from crop use through morphological or genetic variability within most of these longer-lived taxa, therefore the history of edible plant use from Island Southeast Asia, through Near Oceania to Remote Oceania will be assessed in terms of the archaeological contexts from which these plants were identified and the timing of their use. As much as possible archaeobotanical data, whether chance or systematic finds has been incorporated into

20 the review. Through this, a chronology of plant introductions in the Pacific economy will be developed.

Pleistocene: Independent origins for cultivation? Of the suite of crops that now dominate agriculture in Tonga, including a number of aroids and yams, many have now been proven to have been first adopted for use within the terminal Pleistocene in Near Oceania, rather than brought during the mid-Holocene by Austronesian speakers from Island Southeast Asia. This review will therefore begin within New Guinea, discussing evidence for the incorporation of root, tuber and tree crops into subsistence before tracing plant use east to Remote Oceania. It will also be acknowledged that this was not a unilinear movement, and that some of these same crops were also incorporated into pre- agricultural diets further west, indicating that plants were also being carried in the opposite direction.

The earliest evidence for the adoption of root, tuber and tree crops that comprised significant components of the Pacific economy prior to European contact is in the New Guinea highlands. Dioscorea yam starch residues found on stone artefacts and charred Pandanus have been recovered from Joes Garden, Kosipe Mission and dated to over 40,000 BP using Themoluminescence (TL) dating (Summerhayes et al. 2010). Large amounts of archaeobotanical data have also been collected from years of excavations at Kuk Swamp. Within the various phases of occupation at the site, microfossils in the forms of phytoliths, starch and pollen of known edible cultigens have been identified and interpreted as evidence for the beginning of cultivation and manipulation of these taxa (Denham 2007; Denham et al. 2003, 2004; Donohue and Denham 2009; Golson 2007; Wilson 1985). Whether these starch-rich plants were incorporated into an existing highland subsistence system or introduced from lower elevations as part of a new system during the early Holocene remains unproven due to a lack of contemporary data from the New Guinea lowlands (Golson 2007). The Kuk sequence has been divided into several phases, the first two of which have been dated to the Pleistocene and early Holocene.

Phase 1 (10,200-9910 BP) represents the earliest evidence of plant exploitation within the upper Wahgi Valley at the wetland margin (Denham 2007:79). A palaeosurface consisting of pits, runnels, stake and post holes arguably representing wetland management for cultivation of corms and tubers (Denham 2007; Denham et al. 2003, 2004; Hope and Golson 1995) is contemporaneous with stone flakes and grinding stones that contain starch residues of taro (Colocasia esculenta) and a variety of yam (Dioscorea sp.) (Fullagar et al. 2006). Additionally, seed phytoliths of the Eumusa section of bananas, particularly Musa acuminata morphotypes, were extracted from sediments below Phase 1, suggesting the presence of this wild variety of Musaceae within the landscape prior to human occupation (Denham 2007; Denham et al. 2003;

Wilson 1985). Phase 2 (6950-6440 BP) has been argued to represent the first unequivocal evidence for deliberate planting at Kuk Swamp. This comprised of multi-cropping mounded cultivation of Colocasia taro and Dioscorea sp., most likely Dioscorea alata or Dioscorea pentaphylla, with Musa spp. and was confirmed through further residue analysis of artefacts combined with locally elevated frequencies of Musa phytoliths in sediments (Denham 2007; Denham et al. 2003; Fullagar et al. 2006). In total, over 30 edible plants were identified from archaeobotanical and palaeoecological investigations in the upper Wahgi Valley. The early to mid Holocene data collected from excavations at Kuk Swamp have since been interpreted to indicate that Musa acuminata sp. banksii was first domesticated or deliberately cultivated in New Guinea, along with the independent domestication of Colocasia esculenta and Dioscorea alata (Denham 2007; Denham et al. 2003, 2004; Donohue and Denham 2009; Golson 1989, 2007).

Other early archaeobotanical data from Island Southeast Asia, New Guinea and Near Oceania appears to support these arguments for pre-agricultural transport of species, or alternatively that the natural dispersal of the immediate ancestors of crops such as various aroids or Canarium predated human settlement of the region (Yen 1993). Haberle (1994, 1995) identified Colocasia pollen from a sediment core in Lake Wanum in lowland New Guinea at around 9000 BP, but argues that this only identifies the presence of this species within the landscape rather than indicating gardening activity. Similarly, Colocasia and possibly Alocasia taro has been identified within starch, calcium oxylate crystal and cellulosic tissue in residues on stone tools from Kilu Cave in the Solomon Islands from 28,000 BP (Loy et al. 1992), indicating exploitation of these taxa in the Pleistocene. Loy and others (1992:910) argue that the implications of this and other data suggest that the northern Solomons and Australia should be included within the natural distribution of these aroids. Further west in Malaysia, research at Niah Cave has demonstrated that Dioscorea alata and cf. Dioscorea hispida were exploited around c. 40,000 BP through fragments of charred parenchyma and starch granules (Barton and Paz 2007:60-62). Fragments of charred rhizomes of aroids were also found in sediments dated to 23,850-23,020 cal BP, along with starch deriving from the Alocasia (Longiloba complex) or Cyrtosperma merkusii that could also be associated with even older deposits. It is argued that Niah Cave deposits represent a relatively broad spectrum subsistence base that included some toxic plants such as taro which required higher energetic costs in food processing, but proving that people were deliberately manipulating the distribution of favourable plants such as these is another matter (2007:72). It is still therefore assumed that the archaeobotanical record of Niah represents evidence for rainforest foraging of naturally distributed plant taxa. Looking beyond these starch-rich taxa, a large number of other fruit and nut species including candlenut (Aleurites sp.), coconut (Cocos nucifera), Pacific almond or galip nut (Canarium spp.), island

22 lychee (Pometia pinnata), Pandanus spp., and Pangium sp. have been identified from deposits within the Sepik-Ramu region of New Guinea at around 5500BP (Swadling et al. 1991).

Lapita colonisers: Innovation and integration Thus far, this review has been concerned with late-Pleistocene to mid-Holocene plant exploitation in the Pacific, culminating in the earliest evidence for agricultural and arboricultural systems which likely integrated endemic cultigens such as bananas, aroids and yams. During the latter half of the Holocene, it is argued that a new intrusive culture, linguistically and genetically distinct from the indigenous people of Near Oceania, arrived in the West Pacific. These people belonged to the Austronesian-speaking Lapita culture, and were the first to colonise Remote Oceania, bringing with them a number of cultigens that have been labelled by some as the ‘transported landscape’ (Kirch 1984). Whether the origins of the Lapita culture can be traced within Island Southeast Asia, or arose after occupation and interaction within Papua New Guinea is still under debate. The archaeobotanical record indicates that, within Near Oceania at least, the Lapita culture integrated some of these taxa that had already been cultivated in New Guinea into broad spectrum subsistence systems (Gosden 1992; Kirch 1987, 1988, 1989; Matthews and Gosden 1997). These cultigens such as aroids and bananas were then subsequently incorporated these into the suite of crops brought to Remote Oceania after 3000 BP.

Within Papua New Guinea, a number of Lapita sites contain preserved micro- and macrobotanical remains of indigenous and introduced taxa. Two sites are water-logged and anaerobic conditions have preserved macrobotanical remains of fruits and nuts from a range of tree crops. Remains from a mid- to late-Holocene site on Arawe Island are identified as deriving from eight genera and six species including Aleurites sp., Cocos sp., Canarium sp., Cordia sp., Cycas circinalis, Dracontomelon sp., Pandanus spp., and Terminalia sp. (Mathews and Gosden 1997:124). Matthews and Gosden (1997) concede that the botanical remains could be the result of natural beach drift, but point out that the best evidence for human involvement is modification through extraction processes. Extraction can be seen from the fragmentation patterns of the Canarium sp. and Terminalia sp., and charring of the Cycas remains (1997:128). Clearly Canarium continued to play an important role as a food source throughout the Holocene, as further evidence of the exploitation of this genera was found at another water- logged site on Mussau Island (Kirch 1987, 1988, 1989). Over 5000 anaerobically preserved seeds and seed cases representing at last 19 taxa, including Canarium sp., were recovered from Mussau (Kirch 1988:337). These included coconut (Cocos nucifera), Tahitian chestnut (Inocarpus fagifer), Corynocarpus caribbeanus, Dracontomelon dao, vi apple (Spondias dulcis), Pometia pinnata, Pangium edule, and tropical almond (Terminalia catappa). Additionally, a number of Pandanus spp. fruit segments were recovered from contemporaneous deposits and shell peelers and scrapers that were argued to indicate the preparation of starchy 23 tubers and corms such as those of Dioscorea and Colocasia (Kirch 1987:177, Kirch 1988:338). Kirch (1987, 1988, 1989) argued that this data represented the first direct evidence that Lapita communities cultivated a wide variety of tree crops that are still used today in Melanesia, indicating that Lapita culture possessed a full component of arboricultural species at 3200-2800 BP.

Gosden (1992:63) argues that the nature of almost identical plant remains from sites in the Arawe and Mussau Islands indicate that these archaeobotanical records are not representative of formative stages of agricultural development during the Lapita period, but instead suggest well-developed arboricultural systems. Therefore, the transition to vegeculture through integration of local and introduced root crop taxa within subsistence still needs to be pinpointed. More recently, Lentfer and Green (2004) analysed the microbotanical record at the Reber-Rakival Lapita site on Watom Island in PNG, with a focus on the phytolith assemblages from three contexts. The presence of both introduced Eumusa and native Australimusa morphotypes indicate that the Austronesians could have brought bananas with them, and possibly at this same stage used the newly encountered cultivars in conjunction with their own varieties, merging these two streams of banana domestication (Lentfer and Green 2004:85).

As Lapita populations expanded into Remote Oceania, further archaeobotanical evidence indicates the continued integration of roots, tubers and tree crops into colonizing subsistence regimes. In Vanuatu, microfossil evidence from the Lapita-associated sites of Teouma, Vao and Urupiv point to the cultivation of aroids such as Cyrtosperma merkusii, a range of yams including D. esculenta, D. nummularia and D. pentaphylla, and bananas by 3000 BP (Horrocks and Bedford 2004, 2010; Horrocks et al. 2009, 2014). It has been argued by numerous researchers that the brackish conditions in beach back-swamps near many early Lapita settlements would have enabled the initial cultivation of saline-resistant crops such as Cyrtosperma prior to the establishment of more labour-intensive irrigation required for other aroids such as Colocasia or Alocasia (Kirch and Lepofsky 1993; Kirch and Yen 1982; Yen 1973a, 1993). Contained within this argument is the assumption that Lapita populations did not have intensive cultivation techniques, beyond an understanding that these aroids grow best in wet conditions. The late materialization of irrigation within island sequences can be seen to suggest that this technology was not transferred, but rather reinvented over generations or millennia (Kirch and Yen 1982:267). Others such as Spriggs (1990, 2002) instead suggest that the similarities between pond-field, island bed and taro pit wetland cultivation techniques across the Pacific indicate that these were not independent innovations, and therefore derived from prior knowledge and experience within founding populations. Moving east to Fiji, microbotanical data derived from Bourewa on Viti Levu (Horrocks and Nunn 2007) contained evidence for the starch, calcium oxylate crystals and xylem vessels of taro (Colocasia esculenta) and the lesser yam (Dioscorea esculenta) at 3000-2500BP. These identifications would 24 therefore appear to support Spriggs’ (1990, 2002) argument but additional work is required to further establish this link.

Micronesia: A different branch Archaeobotanical research in Micronesia demonstrates the transfer and utilisation of crops east of Island Southeast Asia within the late Holocene. On Kosrae, Athens and others (1996) have collected both macrobotanical and microfossil data upon the antiquity of plant use within archaeological and palaeoecological sequences. Wood charcoal of breadfruit (Artocarpus spp.), Thespesia and Cordyline, along with the charred endocarp and seeds of coconut (Cocos nucifera), Morinda citrifolia, Pandanus, Terminalia and Inocarpus, and storage parenchyma of Alocasia taro and cf. Dioscoreaceae were recovered from excavations at Katem (1996:843). The pollen record from Tafunsak contained abundant giant swamp taro (Cyrtosperma merkusii) pollen coinciding with the earliest period of occupation at 1997-1709 BP, while breadfruit pollen was only present in the latest interval at 1264-1150 BP (1996:843; Pickersgill 2004). This information combined with the micro-charcoal record from the same core indicated that a lowland agroforest was established on the island at a very early date, and further burning was not needed as this would only damage existing crop trees (Athens et al. 1996:843).

Further south, Di Piazza (1998) identified evidence for introduced taxa on Kiribati through analysis of wood charcoal and endocarp within an earth oven dated to around 1430 to 1645 AD. A number of species were interpreted as fuel sources including Cocos nucifera, Cordia subcordata, Pemphis acidula, Guettarda speciosa, and Morinda citrifolia, as well as edible remains such as Pandanus tectorius that had cooked in the oven. The pounding and eating of the drupes of Pandanus have been recorded ethnographically (Di Piazza 1998), as well as the utilization of the uneaten portions as the preferred fuel to ensure fires last long enough to cook pigs or Cyrtosperma. Ethnographic and historic accounts highlight the importance of the giant swamp taro within subsistence throughout Micronesia, as this saline-resistant crop can be grown relatively easily within atoll environments. Although it has recently been discovered that Cyrtosperma is actually endemic to Micronesia (Athens and Stevenson 2001), pollen of this species has also been recorded at 4500 BP within the Ngerchau core in Palau (Athens and Ward 2001) but there is no supporting archaeological evidence for occupation.

Island Melanesia and Polynesia: Evidence for extensive and intensive cultivation It is often assumed that early human arrival and colonisation in Remote Oceania introduced a number of core cultigens within the taro-yam complex, and arboricultural taxa into a large variety of island environments. As the socio-politics and material culture of these colonizing populations evolved into what is sometimes termed Ancestral Polynesian Society (APS) within Western Polynesia and then expanded east to the Society Islands, Tahiti, Marquesas, Easter Island, Hawaii and New Zealand, plant cultivation also evolved to suit local growing conditions.

The difficulty of transporting crops by canoe over large stretches of ocean and plant translocation led to the exclusion of some species from localised subsistence strategies. Post- Lapita archaeobotanical evidence from islands within Remote Oceania has elucidated the timing of these introductions, as well as the contexts within which these taxa were cultivated or exploited.

This section of the review begins with an assemblage of microbotanical remains that are not associated with Ancestral Polynesian Society, but instead with Podtanean post-Lapita deposits in New Caledonia that are synchronous with this cultural transition in Polynesia. Horrocks, Grant-Mackie and Matissoo-Smith (2008) identified cultigens including C. esculenta, D. esculenta and one or more undifferentiated species of Dioscorea from identification of starch granules and calcium oxylate crystals within archaeological deposits dated to 2700-1800 BP in Me` Aure` Cave. The presence of these taxa was interpreted to fill a crucial gap in establishing “a continuous spatio-temporal record of ceramic-age agriculture across the western Pacific (2008:179). Further to this, Dotte-Sarout (2010; Dotte-Sarout et al. 2013) investigated whether arboriculture could be identified from the anthracological record at precolonial Kanak sites in New Caledonia, dated after 1300 AD. Results indicated a continuance of a forest gardening tradition focusing on Ficus sp., Syzygium malaccense, Aleurites moluccana, Hibiscus tiliaceus, Cordia subcordata, Calophyllum inophyllum, Artocarpus altilis, Cocos nucifera, Terminalia catappa among others, originating from the Austronesian/Indo-Pacific sphere of interaction (2013:133).

The Polynesian triangle is bound by the island groups of Hawaii in the north, New Zealand in the south, and Rapanui to the east. This boundary defines the culturally and genetically distinct Polynesians from the Melanesians of Fiji, Vanuatu, New Caledonia and islands further west. The botanical record of Hawaii is important for investigating the expansion of cultivars to the northern extreme of Polynesia. Along with Tonga, the Hawaiian archipelago represents one of only a small number of cases where dryland agriculture was adopted (Yen 1982). It is often presumed that where possible, irrigation for taro will be developed and that dryland techniques are a secondary choice (Spriggs 1982; Yen 1982). Both macro- and microbotanical remains demonstrating the nature and extent of dryland crop utilisation have been recovered from a temporally and geographically diverse range of archaeological sites. Douglas Yen was the first to apply modern identification techniques to assemblages of plant remains from dry rockshelters and open sites on Kaho’lawe Island, Molokai, Oahu and Hawaii Island (Allen 1984). From these sites, a number of primary (Dioscorea alata, Ipomoea batatas, Cocos nucifera, Musa sp., Lagenaria sp, Saccharum sp., Piper methysticum) and secondary cultigens (Canarium sp., Pandanus sp., Cordyline sp., Aleurites moluccana) were identified (1984:22-23). Following this, several projects targeted the recovery of carbonised and desiccated plant remains using systematic flotation and sieving techniques. Rosendahl and Yen 26

(1971) recovered complete and fragmentary carbonised remains of sweet potato (Ipomoea batatas) dated to between 1425 and 1725 AD, while Allen (1983, 1984) identified small fragments of desiccated taro corm along with noni (Morinda citrifolia), coconut and kou (Cordia cf. subcordata) that were interpreted as evidence for purposeful environmental modification. Microfossil evidence from phytoliths and starch has thus far lent support to identification of macrobotanical remains. Musa phytoliths were located within samples taken from the Kona field system, along with the starch and xylem vessels of a root crop cautiously identified as Ipomoea batatas, dated to 1300-1625 AD (Horrocks and Rechtman 2009). In an attempt to combine both macro and microbotanical analyses, Kirch and others (2005) extracted the charred remains of Cordyline fruticosa and many seeds from an earth oven from Kahikinui dated to 460-280 cal BP, along with pollen, phytolith and micro-charcoal assemblages from sediments that were interpreted as evidence for mulching practices consistent with those techniques recorded ethnographically in Hawaii.

To the east, Rapanui, or Easter Island, has been the focus of archaeological projects seeking to answer questions surrounding the enigma of the Moai statues and the prehistoric Polynesian culture that created them. Despite this, very little archaeobotanical research has been initiated. The question of agricultural development within subsistence practices has been raised, and most often dealt with through consideration of the context of dryland agricultural features (Stevenson et al. 1999; Wozniak 1999) and land evaluation (Louwagie and Langohr 2002; Louwagie et al. 2006). The first to address agricultural change through the implementation of archaeobotanical techniques was Cummings (1998) at La Perouse Bay. The contents of a possible lowland garden-pit feature were analysed, and the microbotanical remains of both sweet potato and taro were recovered in the form of pollen and starch granules, respectively. Following this, Horrocks and Wozniak (2008) conducted a microfossil survey of sediments within transects near ahu, house sites and presumed horticultural areas. These deposits were found to contain pollen deriving from bottle gourd (Lagenaria siceraria), and starch of the common yam (Dioscorea alata), sweet potato, and taro. The presence and abundance of these taxa, along with pollen evidence of forest clearance, were interpreted as reflecting the intensive mixed-crop, dryland production of these cultigens, dominated by yams and sweet potato, and supplemented by taro and bottle gourd (2008:16). More recently, Horrocks and others (2012) cored lake sediments in the crater of Rano Kau, and used a combination of light microscopy and Fourier Transform Infrared Spectroscopy (FTIS) to assess microbotanical evidence for agricultural practices within the crater. Some starch granules were much degraded, but were able to be tentatively identified as that of sweet potato, the common yam and taro using FTIS (Horrocks et al. 2012:195-198). In addition, pollen and phytoliths of other cultigens such as paper mulberry (Broussonetia papyrifera), bananas, and possibly bottle gourd were also

27 identified and used to argue that the crater was the location for a mixed-crop horticultural system after 1610-1410 cal BP.

Similar to Rapanui, New Zealand in the southern point of the Polynesian triangle is a case study for archaeological projects investigating dryland subsistence in the lower latitudes, specifically the subantarctic convergence zone of the Pacific. A large array of microbotanical studies have been conducted on Maori gardening soils, gardening features such as stone walls and mounds, house sites, coprolites, and within lake sediment cores from throughout the North and South Islands (Horrocks 2004; Horrocks and Barber 2005; Horrocks and Lawlor 2006; Horrocks, Smith, Nichol, Shane and Jackman 2008; Horrocks, Smith, Nichol and Wallace 2008; Horrocks et al. 2002, 2003, 2004, 2007, 2011). Within these sediments a number of prehistoric introductions are identified from diagnostic starch granules, phytoliths, pollen and xylem vessels. The identified taxa include paper mulberry, sweet potato, Colocasia taro, bottle gourd, and yams, indicating that multi-cropping of selections of these crops was common in production systems that utilised lithic mulching and mounding to enable production of tropical cultigens in a cooler temperate climate. Production was argued to have been supplemented by the exploitation of native taxa such as bracken (Pteridium spp.). Almost all of this data has been collected from deposits that have been radiocarbon or AMS dated to post- 1300 AD, or are above the Kaharoa tephra (dated to 1300 AD). Currently there is no direct evidence for agricultural practices from first settlement (around 1200-1300 AD) until this period.

On the outer edges of the Polynesian triangle, islands such as Futuna, Pitcairn and the Marquesas Islands have also received attention from archaeobotanical research. Piazza and Frimigacci (1991) analysed pollen records from three cores near water sources on Futuna dated from around 600 BP to the present. Within these cores was direct evidence for the giant taro (Alocasia), bananas (Musa spp.), and some species within the family Moraceae argued to most likely be breadfruit (Artocarpus altilis) (Piazza and Frimigacci 1991:131). These crops were interpreted as evidence for the antiquity of pond-field construction and swiddening practices, which could be traced back to the Kele Mea period that pre-dated the development of socio- political stratification, involving the the emergence of strong chiefly titles on the island. Turning to the southeast, Hather and Weisler (2000) and Horrocks and Weisler (2006) have carried out micro- and macrobotanical analyses of sites on Pitcairn Island in the Pitcairn Group. Leaf parenchymatous tissues of Cyrtosperma merkusii were recovered in situ within archaeological deposits associated with prehistoric earth ovens, dated from 1000-1600 AD (Hather and Weisler 2000) Following this, Horrock and Weisler (2006) analysed deposits within a stratigraphic profile from a drainage ditch, and found evidence for the cultivation of sweet potato (Ipomoea batatas) and taro (Colocasia esculenta) through the presence of starch and xylem vessels consistent with these taxa. Unfortunately, there was limited age control in the disturbed profile, and there is a possibility that these remains date to the historic period after the Bounty mutineers 28 settled the island. Work by Allen and Ussher (2013) on the island of Nuku Hiva in the Marquesas has recently identified the presence of sweet potato on shell scrapers dated to 1200- 1400 AD. This represents the earliest direct evidence for the cultivation of sweet potato within the archipelago. A number of other economic cultigens were also identified from residues on multi-use shell tools from occupation sites in the Anaho Valley dated to after 1400 AD, including kava (Piper methysticum), breadfruit, taro and a variety of yam (Allen and Ussher 2013:2810).

So far, this review has investigated archaeobotanical data for plant introductions and cultivation in the outer limits of Polynesia. It will now turn to the interior of the triangle and assess the research that has been carried out in the Cook Islands, Samoa and the Society Islands. Chance finds of macrobotanical remains from Mangaia in the Cook Islands and Upolu in Samoa have been identified by Hather (1994; Hather and Kirch 1991) using a systematic identification system that has enabled the classification of parenchymatous fragments of roots and tubers. On Mangaia, both charred epidermal tissues and unattached epidermal fragments identified as peelings and the edible root sweet potato were collected from the Tangatatau rock-shelter in conjunction with wood charcoal and the charred drupes of Pandanus tectorius (Hather and Kirch 1991). The research was significant as it suggested that the sweet potato was in central eastern Polynesia by 1000 AD, and was argued by the Hather and Kirch to lend credence to Yen’s tripartite hypothesis for the distribution of this cultigen in the Pacific (1991:892-3). The tripartite hypothesis proposes three pathways for sweet potato into Oceania: a prehistoric transfer from South America to Polynesia, transport by the Spanish from Mexico to the Philippines, and a later introduction from Europe to the East Indies and New Guinea (Montenegro et al. 2008).

Further east, parenchymatous remains of Dioscorea spp. were identified using Scanning Electron Microscopy (SEM) from excavations carried out by Green and Davidson (1969, 1974) on the island of Upolu in Samoa (Hather 1992, 1994b). The fragments were 2-6mm in size and were identified to genus from the morphology of vascular tissues, but the state of preservation and lack of epidermal tissues inhibited classification to species. More recently, Kahn and Ragone (2013) identified fragments of breadfruit (Artocarpus altilis) exocarp on house platforms in the ‘Opunohu Valley on Moorea, Society Islands dated to around the middle of the 17th century. This built upon the chronology for the cultivation of breadfruit in Tahiti established by Orliac (1997) when charred breadfruit wood was recovered an earth oven from the Papeno’o Valley in rock-shelter deposits dated to the 14th century.

Tonga: The gap in the archaeobotanical record Within Western Polynesia, the Tongan Archipelago represents an important location for archaeological research. This island group is acknowledged both as the homeland for the

29 colonisation of East Polynesia, as well as the terminus for Lapita migration. Despite this, only a very small number of palaeoecological studies have contributed information upon Polynesian introductions and associated cultivation practices in Tonga. Fall (2005, 2010) and Fall and Drezner (2011, 2013) document the first evidence of Colocasia in pollen cores taken from the Avai’o’vuna Swamp and the Ngofe Marsh within the Vava’u group, and on Eua in the Tongatapu group from 2600 BP. Other introductions at this time included Casuarina equisetifolia, Cordyline fruticosa, and Pometia pinnata. Indigenous species such as Canarium harveyi, Cocos nucifera and Pandanus tectorius display an increase in pollen associated with Lapita settlement (2900-2600 BP) that may indicate their cultivation for food. In addition to these Lapita-associated introductions, Fall (2010) documents the introduction of the Polynesian cultigen Ipomoea batatas as a historic introduction into the archipelago. This supports Kirch’s (1978, 1990) argument that the adoption of this versatile South American cultivar in Tonga coincided with the arrival of Europeans. However, the cultivation of sweet potato was documented in Tonga during Cook’s second voyage in 1777, which may in fact suggest prehistoric rather than historic transfer of this cultigen. No macrobotanical remains have been recovered from archaeological sites within the archipelago that could corroborate these identifications, and there is a significant lack of horticultural features in the archaeological record. The absence of an archaeobotanical record for Tonga is significant due to the importance of these islands for Pacific migration, and therefore a targeted and systematic regime of flotation, wet-sieving and bulk soil sampling for macro- and microbotanical remains the logical approach to address this problem.

Tonga in the Pacific: A summary Around the Pacific, archaeologists and palaeoecologists have begun to use multi-disciplinary approaches that combine natural and cultural datasets to enable the development of chronologies for localised and regional plant introductions and utilization. From the identification of taxa within the archaeobotanical record, interpretations have been made about the movement of crops around the Pacific region through island colonisation and trade, as well as the nature of crop production through localised innovation and adaptation. It appears that the taro-yam complex was the most extensive horticultural system and was transported from Near Oceania into Remote Oceania by the Lapita cultural complex to all of the outer limits of East Polynesia.

Current archaeobotanical data reflect the movement of human and plant populations throughout the Pacific, but also local environmental limitations and cultural practices that affected the number and types of cultigens that were grown. Further east, islands become higher and entirely volcanic in origin, making irrigation ineffective in more temperate climates. It is reasonable to assume that these island populations relied less on those aroids that required some form of irrigation, such as Alocasia and Cyrtosperma to ‘intensively’ cultivate, and instead ‘extensively’ cultivated dryland crops through mounding such as yams (Dioscorea sp.) and 30 sweet potato (Ipomoea batatas) or the arboricultural production of breadfruit (Artocarpus sp.) and bananas (Musa sp.). However, questions have been raised regarding how to define intensive agriculture, and how visible these cultivation techniques are within the archaeological record (Leach 1999).

The assumption that population growth and social production are the primary drivers for intensification, begs the question as to how to explain cases such as the Tu’i Tonga chiefdom under current typologies where intensification in the traditional sense (Boserup 1965) is defined as increased labour input holding land as a constant. By European arrival, almost all of Tongatapu was dedicated to agriculture, but to fit this classic definition for intensification this dryland production would be required to have developed from swidden agriculture with a shortening of fallow length in plots, followed by permanent plot boundaries and a trend towards rain water control and soil additives (Yen 1973a). A chronology of horticultural production is required to identify this process, something currently lacking within archaeological research in Tonga. In light of this, this thesis investigates the development of production systems over time using Tongatapu as a case study, and seeks to elaborate upon and test modes of production other than the classically defined intensive techniques, using archaeobotanical data and agroecological modelling.

PART ONE- AN ARCHAEOBOTANICAL COMPARATIVE COLLECTION FOR TONGA

Chapter 3 Reviewing Microbotanical Analysis

This study uses a combination of both micro- and macrobotanical techniques to research subsistence strategies adopted within the Tongan archipelago in prehistory. Starch analysis and the study of charred parenchyma are both areas of archaeobotany that have been under-utilised due to perceived issues with taphonomy, extraction and the diagnostic value of morphological characteristics of these botanical remains. This chapter explores past research, and demonstrates that the cumulative data from experimental studies, ethnography and applications of these findings within archaeological research has enabled a greater understanding of human-plant interaction. The current study will utilise the results of past research into taphonomy and identification techniques, and develop these further to enable later classification of these micro- and macrobotanical remains extracted from sites on Tongatapu within Part Two of this thesis.

Although it is a relatively new area of archaeobotany, starch analysis has been applied to answer a range of archaeological questions regarding population migration, diet and subsistence, tool function, and human-environmental interaction. Yet the chemical and biological properties of starch as a carbohydrate, and how these can change under different conditions such as heating or cooling, are still mostly not well understood. Reichert (1913) conducted the first scientific study of starch and its properties, and his seminal guide is often used in microbotanical research. Variation in starch morphology at different taxonomic levels is well known, and has been used to identify starch found in sediments (Horrocks et al. 2004, 2008; Horrocks and Lawlor 2006; Horrocks and Rechtman 2009; Lentfer et al. 2002; Therin 1994; Therin et al. 1999) or as residues on stone artefacts (Barton 2007; Field et al. 2009; Fullagar 1998, 2006; Fullagar et al. 1998, 2006; Loy et al. 1992; Loy 1994; Pearsall et al. 2004), pottery (Crowther 2001, 2005, 2009), and dental calculus (Boyadjian et al. 2007; Henry and Piperno 2008; Reinhard et al. 2001).

Biology of starch and identification potential The biological characteristics of starch grains offer a unique picture of plant use in the past. Starch is produced as a form of energy storage during photosynthesis and is found in plastids in every kind of tissue in most green plants. Light energy is converted to potential energy within the chloroplasts found in leaves through the splitting of water into hydrogen and oxygen (Banks and Greenwood 1975; Gott et al. 2006). The freed hydrogen combines with carbon dioxide from the atmosphere to form glucose, which is stored in two forms. It can be hydrolysed, synthesised and then stored overnight as reserve starch within the amyloplasts in storage organs such as roots, tubers, fruits and seeds (Cortella and Pochettino 1994:172; Jane et al. 1994), or used during the day as transitory starch in chloroplasts (Banks and Greenwood 1975:1). Transitory

33 starch is much smaller and less diagnostic than reserve starch and is mostly found in the leaves, stem and petiole (or leaf stem) (Banks and Greenwood 1975:1; Gott et al. 2006; Messner 2011).

Starch itself is biologically defined as discrete insoluble semi-crystalline polymers consisting of linear amylase and highly branched amylopectin. The internal structure of these granules is composed of alternating semi-crystalline and amorphous growth rings, and an amorphous core (Cai and Wei 2013). Granules are formed within the plastids from a nucleation point, or hilum, from which these layers of amylose and amylopectin are accreted (Loy 1994:89). The optical pattern created by these layers is called ‘lamellae’, and these increase in density and insolubility towards the outer margins of the granule. Three different types of crystalline structures have been identified within starch granules through X-ray diffraction techniques, designated as A-, B- and C-types (Banks and Greenwood 1975:242; Gott et al. 2006). A-types contain a pattern that is visible in most cereals, while B-types are seen commonly in tubers and C-types are within most root and seed starches (Cortella and Pochettino 1992:178). Heat and moisture during plant growth dictate the percentages of each type produced within the plastids.

Partly due to the semi-crystalline structure of starch, granules are also birefringent in polarised light, displaying a ‘Maltese’ or ‘extinction cross’ pattern. This effect is created by light passing through the grain which is deflected owing to the differences in density and structure of the various lamellae. This deflection creates interference which in turn results in the generation of localised bright and dark regions that extend radially when polarised due to the spherical shape of granules (Loy 1994:89). These interference patterns are not unique in their physicochemical structure and can sometimes be confused with synthetic semi-crystalline polymers which form spherulitic structures that are not unlike starch in the early stages of growth when viewed in polarised light (Banks and Greenwood 1975:247). However, it is the combination of attributes that are visible in both polarised and brightfield light that permit the direct identification of starch grains microscopically.

The growth of starch within the plastids is under genetic control, and size and shape of granules is determined by the conditions within the cell. The type of starch granule that grows is also dictated by the plastid, where a single nucleus gives rise to a ‘simple’ starch granule and a multiplicity of nuclei form ‘compound’ granules (Banks and Greenwood 1975:251; Gott et al. 2006:41; Loy 1994:90). Each granule within a compound granule exhibits its own birefringence pattern and has the potential to separate and become simple granules. Other traits such as the morphology of the hilum region including vacuoles and fissuring, and the density, hydrophobicity (insolubility) and thickness of the accretion layers are also inherited genetic effects (Loy 1994:91). Because of genetic control over starch morphology, there is the potential to identify the botanical origin of native starch. Some granules within a population are highly

34 diagnostic, whereas others are less useful for identifying the species of origin due to features that overlap with other taxa (Gott et al. 2006:40). It is these diagnostic or ‘signature’ types that are mainly used to identify unknown taxa found in archaeological assemblages.

Many studies of species-specific starch morphology and physicochemistry have been conducted to assess the range of variation within any given sample at different taxonomic levels. Initial studies only used light microscopy (Gott et al. 2006; Barton and Fullagar 2006; Reichert 1913) but more recent use of Scanning Electron Microscopy (SEM) enables high resolution imaging of the three-dimensional shape of starch (Barton and Fullagar 2006:52; Jane et al. 1994; Lindeboom et al. 2004). Comprehensive analyses of starch morphology have demonstrated that attributes such as the presence of lamellae, hilum features, three- and two- dimensional shape, shape modifiers such as faceting, and overall size, characterise different biological taxa. Further, there is also evidence that starch can vary morphologically within different organs of the same plant (Gott et al.2006; Jane et al. 1994:46; Loy 1994; Reichert 1913), which suggests that if these features were also diagnostic, it could be particularly useful to archaeologists attempting to understand plant exploitation and processing in the past.

Starch taphonomy Detailed studies in both archaeobotany and the food sciences have emphasised the range of changes that can occur when native starch is exposed to a range of physical, chemical and biological environments. A number of methodological studies have focused on these taphonomic issues, highlighting the changes that can affect native starch biochemistry and morphology under differing conditions both prior to and after deposition, and how these factors can influence the identification and quantification process (Babot 2003; Crowther 2009, 2012; Haslam 2004; Henry et al. 2009; Weston 2009).

Plant processing Archaeobotanical research targets plant remains that are found within archaeological, and therefore anthropogenic, contexts. Most starch remains found within domestic deposits such as occupation floors or middens derive from economic or supplementary plant taxa that have been processed in some way. Roasting, charring, boiling, milling, pounding, freezing, dehydrating, and rehydrating are all examples of cultural food processing techniques that can affect the morphology and chemistry of starch granules (Babot 2003; Crowther 2009; Henry et al. 2009; Laurence 2013:34-36; Messner and Schindler 2010). The effects of various cooking techniques on starch have been by far the most researched of these processing techniques. Experiments have been tailored to assess the range of the conditions under which starch either melts or gelatinises when exposed to water and/or heat.

The thermal conversion of starch is defined when “…starch is converted from an ordered, semi-crystalline to a disordered, amorphous state during cooking…” (Crowther 2012, 35

2009:23). Gelatinisation can occur when starch is exposed to both moisture and increased temperature, causing swelling and a loss of birefringence (Banks and Greenwood 1975:259-67; Messner and Schindler 2010). Species-specific water-dependent temperature thresholds affect the nature and timing of starch thermal conversion (Crowther 2009; Henry et al. 2009:917), but complete and uninhibited gelatinisation of all granules can only occur when more than 60-65% water is present. Within most Pacific cultigens the temperature range for gelatinisation under excess moisture is around 60-85˚C (Moorthy 2002). Partial gelatinisation is possible when these conditions are not met and the amylopectin crystallites only partly melt, resulting in an incomplete loss of birefringence and limited morphological change. Swelling is also reversible up to a point when loss of optical properties begins, which occurs within a population of starch grains over a relatively narrow temperature range of 5-10˚C (Banks and Greenwood 1975:260).

Melting of starch crystallites occurs when granules are dry-heated, through charring or roasting at temperatures above 220˚C. Starch heated in the absence of moisture will thermally degrade through breaking down into smaller glucose units and eventually carbonise before becoming structurally disordered (Crowther 2012, 2009:26). Experimentation has demonstrated the visible changes that can occur during these types of cooking techniques. Babot (2003) roasted samples of corn (Zea mays L.) kernels and quinoa (Chenopodium quinoa Willd.), reporting that these granules displayed various combinations of flat relief, weak birefringence, deformation of the extinction cross, clumping and slight gelatinisation, but not all grains showed gelatinisation features. Babot concluded that the water content at time of cooking and the heating temperature were responsible for the damage (Babot 2003:73). Similarly, the effects of charring were evident in changes in granule birefringence, swelling and clumping, but were dependent on grain size (Babot 2003:74). Starch size is related to hydration, where water molecules inhabit the crystalline regions of the amyolopectin molecules (Torrence and Barton 2006). Smaller starch granules have less capacity for swelling, while gelatinisation occurs more quickly for larger granules unless these granules have a high ratio of amylase to amylopectin which buffers gelatinisation (Banks and Greenwood 1975:260; Crowther 2012; Saul et al. 2012:3484).

Studies also demonstrated that gelatinization is not as simple as dry heat versus wet heat cooking, or charring and roasting versus boiling. Consideration must be given to the role that heat-transfer mechanisms and micro-climates have in the cooking process (Messner and Schindler 2010). In earth ovens the thermal agent is heated rock. When these are placed in direct contact with starchy organs, the oven is instantaneously heated through trapping hot moist air which emanates from the food and is then vaporised when this comes in contact with the rocks, resulting in complete breakdown of starch within the oven (2010:334). In contrast, when the external heat source is in the form of fire on top of the earth oven that increases in temperature gradually, the moisture is slowly evaporated from the organ and starch is damaged but can still 36 survive cooking. Messner and Schindler (2010) therefore argue that “…the environmental conditions generated by the heat transfer systems were of greater influence to starch degradation than temperature alone.” Different water-uptake mechanisms in different wet cooking methods also affect the rate and nature of gelatinization. For example, boiling increases water absorption at a faster rate, but steaming allows more uniform absorption over time (Crowther 2009:28-29). These factors, and the type and order of cooking procedures followed can greatly affect the archaeological visibility of particular activities. Two of the main methods for cooking starchy foods in the Pacific throughout prehistory have been by steaming in earth ovens and boiling, so these studies’ findings are particularly relevant to this study.

Other studies have demonstrated that the temperature during charring can significantly alter the nature of starch damage. Low temperatures up to 220ºC will not commonly cause any morphological changes, but higher temperatures up to 350ºC can cause complete loss of structure or fusion of granules similar to those observed within boiled samples at lower temperatures (Valamoti 2008). Similarly the location of starch within an organ that has been cooked whole at any temperature will limit the amount of damage. For example, granules in the centre of grains are more likely to survive any cooking method as these are protected from the effects of liquid and heat (2008:269). Some alteration of morphology, such as slight swelling, is still possible.

Crowther (2012, 2009) reviews the potential for starch survival within charred and carbonised residues on ceramics, and argues that these are ideal contexts for recovering non- gelatinised granules as evidence of cooked foods under particular charring conditions. The temperature must not exceed starch thermal degradation temperature, but this depends on the intensity of the heat source, the distance of the vessel from that source and the location on the vessel where the granules are located. Secondly, desiccation and carbonisation must occur relatively quickly before all starch granules undergo gelatinisation. Starch granules of underground storage organs (USO) such as roots and tubers are less likely to survive charring without gelatinisation due to the high water content within these organs, biasing the archaeological record towards cereals and legumes where these organs are utilised (2012).

Further studies have looked beyond the morphological changes that can occur within starch granules exposed to charring, and instead attempted to assess whether the survival of starch granules within charred residues reflect the intensity of plant use. Raviele (2011) replicated charred maize residue construction under controlled conditions, where the variables tested were the ratio of plant extract to water and meat, and the type of maize (Zea mays) (whole green cob, whole green kernel, lightly pounded dried kernel, whole dried kernel and ground maize flour).The overall results did not demonstrate a trend for increasing abundance with increasing proportions of maize in residues, suggesting differential survival of starch during

37 charring. Despite this, it was noted that dried kernels and cob had higher abundances of starch survival, most likely a result of high starch production within maize at this time of organ development (Raviele 2011:2711). Similarly, Saul and colleagues (2012) experimentally replicated charring conditions for einkorn (T. monococcum) and acorn (Quercus sp.) in ceramic pots at more than 100ºC for three hours. These experiments indicated that starch can survive repeated charring episodes in relatively high quantities (184 granules mg-1 and 608 mg-1 respectively).

Other food processing techniques such as parching, fermentation, freezing and milling can also cause distinctive damage to native starch granules. Henry and others (2009) tested different cooking techniques on both ground and whole legumes and grains. One of these tests involved parching three millilitres of each sample for three minutes in a muffle furnace at 200ºC. The authors noted a significant degree of heterogeneity within the sample, but argued that parching caused the most distinctive damage to granules (Henry et al. 2009:918). Most granules appeared encrusted with small particles that were either small starch granules or other organic material, which does not occur through any other cooking technique. Other common morphological changes associated with parching were the definition of lamellae and development of deep radial fissures (Henry et al. 2009:921). Within the same study, starch granules were exposed to yeast to replicate fermentation. Whole samples displayed surprisingly little change except for a possible extra ‘arm’ within the extinction cross; however, granules within the ground samples often displayed signs of ‘hollowing out’ at the hilum. (2009:921).

Effects of freezing on starch granules from several cereals, tubers and legumes has also been tested, and indicate that this process changes both the chemical properties of the starch granules, specifically a loss of birefringence, and also the physical appearance of the grains in terms of flat relief, fragmenting, fissuring, breaking and bursting (Babot in Beck and Torrence 2006:66-67; Babot 2003:74). Milling involves the application of friction to plant parts and their by-products, and is a technique most commonly used for separating the starchy endosperm of grains from the pericarp (Henry et al. 2009:916). Experimentation has revealed the extensive damage that milling can have on starch granules, which includes truncation, incompleteness, fracturing, collapsing, and bursting (Babot 2003:76). Some damage also can occur at the hila, with fissuring and large open vacuoles. In contrast, short term pounding using stone implements has been demonstrated to have very limited affect on starch granules aside from the disaggregation of compound granules (Robertson, in Beck and Torrence 2006:68-69).

Damage caused by these various food processing techniques can render starch unrecognisable as such in archaeological contexts. Studies have therefore attempted to use other chemical tests to identify the presence of cooked starch residues on artefacts or in sediments.

Congo Red (empirical formula C32H22N6O6S2Na2) stain has been used in the agricultural and

38 food sciences as a contrast stain for cellulose, starch and amyloid fibrils, but the potential of this stain to test for cooked or damaged starch in archaeological contexts has only been recognised within the last two decades (Cortella and Pochettino 1994; Lamb and Loy 2005; Lamb 2003; Reichert 1913; Weston 2009). The stain only binds to proteins in alkaline or acid buffer conditions, so at a neutral pH only starch and cellulose will be stained. Unaltered or undamaged starch is hydrophobic and so does not take up the stain, but starch that has become disordered allows Congo Red to react with the amylase content of these granules resulting in a red discolouration with an orange-red or green-gold glow in cross-polarised light (Lamb and Loy 2005:1434). Trypan blue has been used in much the same way, penetrating the outer layer of damaged starch granules to stain the exposed interior blue (Barton 2007:1734).

Some researchers have argued that these stains are not reliable indicators of the presence of starch within archaeological contexts, and have subsequently experimented with thermally stable α-amylase, an enzyme that degrades the chemical linkages contained within starch (Hardy et al. 2009). These tests involve exposing a sample of extracted residue to α- amylase and observing the degradation of objects tentatively identified as starch (2009:251). Alternatively a sub-sample can be used as a control by adding α-amylase, and comparing this to the remaining analysed portion of extracted residue (Saul et al. 2012). If starch is thought to be present in the remaining sample but not visible within the control, then this is a good indication that the identification of starch within an archaeological sample is correct. However, this can be problematic as it is not possible to confirm the identification of individual granules.

Starch in sediments Soil properties such as pH levels, temperature, texture and moisture content, in conjunction with soil constituents including enzymes, bacteria, fungi and earthworms, play the greatest role in influencing starch degradation and movement after deposition (Barton and Matthews 2006; Haslam 2004:1721). These factors are particularly relevant in tropical climates such as Tonga which generally have low rates of organic preservation (Hather 1994). A large array of studies has been carried out under varying soil conditions, either as individual variables or in conjunction with one another. These experimental studies been utilised by archaeologists and archaeobotanists to further our understanding of the extent to which morphological and chemical changes can occur after deposition within archaeological contexts.

One of the most common causes of starch degradation within tropical sediments is through hydrolysis, whereby starch is chemically broken down by polysaccharidases (enzymes) produced by bacteria and fungi. When plant and animal cellular material decays, these enzymes are released into the soil in significant quantities and are present in almost every soil type encountered throughout the world (Cheshire et al. 1974). Two classes of enzyme-producing bacteria are recognised: autochthonous, which grow slowly and predominate when there is little

39 oxidisable substrate and zymogenous, which respond to substrate addition by rapidly increasing, with the majority dying out after substrate exhaustion (Burns 1982; Haslam 2004). It is therefore possible for inactive enzymes and the bacteria that produce them to exist in the soil prior to starch deposition. The presence of these enzymes, and their potential to remain inactive in the soil substrate, limits the potential for starch survival in archaeological contexts.

To gauge the extent and rate of starch degradation, several studies have replicated hydrolysis in vitro (Mellon et al. 2002; Steup et al. 1983). Results indicate that the majority of both transitory and reserve starch is degraded within the first few days of exposure to enzymatic digestion, with rates of decomposition after this following an asymptotic curve (Haslam 2004:1720, Barton 2009). Within this generalised trend there are species-specific differences in enzyme attack patterns and rates, depending upon factors such as granule structure and size, amylose to amylopectin ratios and crystal types (Haslam 2004: 1720; MacGregor 1980; Zhang et al. 2005). Haslam (2004) compiled these studies into a table that archaeobotanists can use to understand and interpret possible biases in the archaeological record. When ranked, taxonomic patterning suggests that the rate of enzymatic degradation increased when granule size and amylose content decreased. Some species such as bananas (Musa and Eumusa spp.) are resistant to enzymatic degradation unless gelatinised (Zhang et al. 2005:144). Once gelatinisation begins the rate of degradation increases as enzymes have easier access to the more susceptible amorphous cores of the granules of most taxa. The temperature at which gelatinisation occurs does not appear to be a significant factor affecting the rate of starch degradation for each taxa. However, as discussed previously, the amount of moisture in the soil regulates the extent of swelling and gelatinisation and thus has more impact (Haslam 2004). One variable that does seem to affect the presence of microbial, enzyme and fungal activity in soils is depth. Enzymatic degradation of starch is much more likely to occur closer to the surface than at midpoints or near basal deposits, where activity is correlated with the presence of other organic matter (Haslam 2004:1721; Taylor and Belton. 2002). The rate of starch degradation is also seen on artefacts, where starch abundance at 108 weeks is already representative of those in archaeological contexts (Barton 2009:135).

Other taphonomic factors that affect starch preservation in sediments include temperature, pH and moisture content of the sedimentary matrix. These variables have the potential to damage starch through causing gelatinisation or damaging the biochemical properties of granules; however, these rarely directly affect starch individually, and so should be considered in conjunction with enzymatic hydrolysis (Haslam 2004). It is the interplay between these variables and the presence of microbial and enzyme activity that cause starch degradation. As mentioned earlier, temperature and moisture cause swelling and shrinking of starch, enabling various enzymes access to the interior of the granule. The same principles described in relation to heating and cooling within food processing techniques is applicable to soil conditions. 40

Hydrolysis can also occur in both acidic and alkaline soils, but is slower than enzymatic decomposition (2004:1721). Some enzymes often favour different pH levels, such as invertase which is more active in more alkaline soils, whereas amylase can affect starch in any conditions (Dick et al. 2000).

Quantitative studies of starch preservation (Barton 2009; Haslam 2009; Therin 1998) have made observations of displacement of starch within sediments over time. Therin (1998) attempted to replicate the downward displacement of starch grains in various sandy matrices using groundwater. Variables tested included sediment particle size, irrigation rate and starch grain size, monitored over a two-month period. The results indicated that there is limited downward movement of starch grains, but within this overall trend patterning related to granule and particle size. Larger starch grains have less chance of becoming mobile, but if they do move, these granules move slowly and have less chance of becoming trapped. Smaller sand particles also correlate with less starch mobility, but an increase in irrigation rate will increase starch movement. Haslam (2009) attempted to take this experimentation a step further, and investigate both upward and downward movement of starch, as well as lateral movement. These parameters mirrored those used by Therin (1998), but only one sand matrix was tested with larger particles of 250-500µm. The sediment was autoclaved for 30 mins at 120ºC to eliminate fungi and bacteria then placed into a PVC pipe ‘cross’ with a small amount of starch (0.1g) in the centre, and placed upright (Haslam 2009:95). The experiment was irrigated at a rate of 160ml twice a day from the top of the set-up for a period of 30 days. The sediment was then sampled at 27 locations within the cross and processed for starch extraction, revealing that around 16% of starch moved downward up to 6cm although the highest number of starch was still found at the centre (2009:98). Very small numbers of starch also moved laterally up to 12cm, and upwards 2cm from the centre. The implications of these experiments are that researchers need to assess factors such as sediment compaction, particle size and local rainfall when interpreting the potential movement of starch in sediments, along with other taphonomic issues such as bioturbation and human activities.

Researchers have sought to provide solutions to taphonomic issues through investigating the types of conditions that are favourable for starch preservation (Barton 2009; Haslam 2009, 2004; Laurence 2013). The presence of heavy metals within soils or high clay content can neutralise enzymes and thus limit the effect that bacteria and fungi can have on starch granules within these deposits (Haslam 2004). Additionally, soil aggregates, or Particulate Organic Matters (POMs), and residues on artefacts can form a protective barrier for starch through limiting surface area accessible for enzyme digestion. Artefact surfaces can also provide a micro-environment that protects starch from the effects of temperature, moisture and downward displacement through groundwater. Studies of starch residues on bone, wood, lithic and ceramic artefacts indicate that the preservation of these residues in soils is possible. 41

However, experiments that attempt to replicate archaeological deposition of tools used for food processing have produced no conclusive results (Barton 2009; Haslam 2004; Lu 2003). The results of one study indicate that starch is in fact more likely to survive as residues on artefacts if the tool remains on the surface for some time, as sunlight can reduce enzymatic degradation (Barton 2009). Another experiment found that starch residues on buried tools had high survival rates of around 75% after 71 days (Lu 2003:124), but the highest quantities of starch were found on tools that were in sheltered conditions on the surface which supports Barton’s (2009) hypothesis.

Modern starch contamination Contamination has plagued archaeobotanical studies involving the analysis of microbotanical remains. Many studies have attempted to understand the nature and sources of contamination that can occur both in situ and during post-excavation processing in the laboratory. Contamination is defined here as starch that can be added to samples through aeolian (airborne) processes or through transmission by direct contact. Laurence (2013; Laurence et al. 2011) provides an extensive review of modern airborne starch contamination that derives from food processing factories such as flour and maize mills, as well as starch that is contained in pollen. Pollen starch is undistinguishable from reserve starch and can be released when pollen ruptures either on the ground or mid-air (Laurence et al. 2011:215). Both insect and wind-pollinated species can produce pollen starch which provides energy for the pollen tube (Baker and Baker 1979). Microscopic slides placed close to sites in Texas where earth ovens were being assessed for ancient starch contained quantities of starch after 96 hours.

Other researchers have recommended that archaeological studies should incorporate an assessment of environmental airborne starch contamination in the field (Loy and Barton 2006; Messner 2011). Yet others recommend sampling sediments surrounding artefacts to assess whether residues on tools are accurate representations of use or contamination (Barton 2007, Loy et al. 1992; Williamson 2006). This is determined through the taxa and quantities present in either sample. This is problematic for a number of reasons. Fullagar and others (1998:51) argue that the: “...difficulty in comparing starch grains on tools with starch grains in sediments is the arbitrary choice of a quantity of sediment for a comparison with a given quantity of residue”. Similarly, as pointed out earlier, starches in sediments can be more prone to enzymatic degradation, so false negative results can be the outcome of these studies (Laurence et al. 2011; Zarillo and Kooyman 2006). It is also difficult to argue that starch found in sediments is contamination or vice versa, if consideration is given to the fact that an area may have been used for food processing and that a tool was left in the same location after use.

Transmission of starch through handling and equipment is another source of contamination in the field. This has been addressed in a number of studies, particularly those

42 concerned with residues on artefacts. Handling of an artefact both during original use and post- excavation processes can cause starch to become attached to both working surfaces and other areas of the tool that are not directly related to use (Barton 2007; Loy et al. 1992). Modern handling contamination can be avoided by wearing starch-free gloves when bagging artefacts into new zip lock bags in the field (Barton et al. 1998:1233; Hart 2011), but often analyses include artefacts that have been excavated without these protocols and so researchers have sought a means to test for levels of contamination. Suggestions to control for contamination include testing various locations on the artefact, proposing that use-related starch is concentrated around working edges (Barton et al. 1998; Loy et al. 1992; Piperno and Holst 1998). Another control is to compare artefact use-wear with the results of starch analysis (Allen and Ussher 2013; Barton 2007; Ussher 2009). Where these are complementary, this provides strong support for food processing technology.

Airborne native starch can not only contaminate sediments in situ or during fieldwork, but also within the laboratory along with starch found in chemicals or through manual transmission. There have been a number of published studies highlighting potential sources of starch contamination in laboratory environments (Crowther et al. 2014; Laurence 2013; Laurence et al. 2011; Loy and Barton 2006; Loy et al. 1992; Messner 2011). The most extensive of these is that carried out by Crowther and others (2014), where not only the consumables and equipment in the laboratory were tested for the presence of starch, but also surfaces and air within various working spaces. This was designed as a comparative study between ancient starch laboratories at the University of Calgary and Oxford University, and the results strongly suggest that starch contamination is a major concern that needs to be addressed before interpreting archaeological material. Consumables were tested by sampling liquids or exterior swabs, while environmental samples were taken using horizontal ‘passive’ traps, or vertical stationary or mobile adhesive traps. High numbers of wheat (Triticum sp.), maize (Zea mays) and potato (Solanum tuberosum) starch were found on particular brands of powder-free gloves, pipette tips, paper towels, and within Calgon and Sodium polytungstate commonly used for heavy liquid separation (Crowther et al. 2014:86). Spaces that had significant quantities of starch included fume hoods, floors and work benches, but these numbers were lessened after cleaning.

Crowther and others’ (2014) study corroborate data from previous studies. Powder-free gloves have been also been tested for starch contamination by Laurence (2013; Laurence et al. 2011) and others within the medical sciences (Campbell et al. 1984; Makela et al. 1997; Newsom and Shaw 1997; Wilson and Garach 1981), and found to have varying quantities of maize starch. These gloves are manufactured in the same factories as powdered gloves, and therefore can easily be subjected to contamination by airborne starch if care is not taken. Similarly, Newsom and Shaw (1997) conducted a survey of airborne starch within a hospital 43 environment and found that an average of 13.8 granules/30L air was present even in areas such as intensive care units. Many of the starch granules observed within these studies displayed signs of morphological and physicochemical modification such as cracking and gelatinisation (Crowther et al. 2014:101; Laurence et al. 2011:228), indicating that the condition of starch cannot necessarily be taken as an indicator of its age.

Protocols that limit the potential for modern starch contamination in the field and in the laboratory have been developed in response to these studies. There are no alternatives for powder-free gloves, but some brands have little or no starch contamination and should be selected over others (Crowther et al. 2014). It is also possible to limit the amount of contact gloves have with samples by using equipment such as sterilised forceps. Limiting or excluding the use of paper products can also reduce the amount of airborne starch within the laboratory environment. All centrifuge tubes, mixing rods, Petri dishes, sieves and pipette tips should be treated for decontamination with 5% sodium hydroxide (Crowther et al. 2014:102) or hypochlorite (Laurence 2013: 52; Laurence et al. 2011) prior to use. These protocols are also replicable in the field, where particular powder-free gloves should be worn when handling samples, and contact with the sample should be limited before placing in zip lock bags. All sampling equipment at the very least should be cleaned with boiling water before use, but disposable items should be utilised where possible. Food should also not be consumed on site (Loy and Barton 2006:165).

Spatial differentiation for various tasks within the laboratory is also essential. Any processing of modern starches for reference collections or experimentation should be carried out in a different room to that where post-excavation processing of sediments is usually undertaken (Crowther et al. 2014; Loy et al. 1992). This is especially important if samples are being dehydrated and then milled or pounded. Consumable and environmental contamination should be monitored on a regular basis to ensure the risk of contamination is low during different stages within the working week or seasons. Monitoring of airborne starch in the field is also a useful measure to at least provide a gauge of the most common modern morphotypes and quantities of starch within the local environment (Crowther et al. 2014; Laurence et al. 2011; Loy and Barton 2006).

Sampling strategies and extraction techniques Research on sampling and extraction techniques has continually forced revision of the methods used to extract starch. These differ technically when researchers are dealing with residues on artefacts or in sediments, but issues of scale influence both. By nature micro-analyses such as microbotanical studies represent high resolution palimpsests of events occurring in the past, but it can be difficult to interpret the spatial and temporal scale of these events. It is therefore essential that the choice of a sampling strategy is related to a specific research question, and the

44 connection between the microbotanical remains and the processes that created the archaeological record is made explicit by comparing samples. It is this comparison that ensures that a sample represents more than just micro-scale data. Here, discussion focuses on the sampling and extraction of sediments, as this relates to the methodological approach employed on Tongatapu in the current research.

Many microbotanical studies target multiple microfossils, and an appropriate sampling strategy incorporates taking samples from a combination of archaeological features and background environmental deposits in swamps or trenches. Samples need to be of an appropriate size, or multiple samples should be taken from the same contexts, due to problems stemming from the use of combined multiple microfossil extraction techniques (Coil et al. 2003; Korstanje 2003; Torrence 2006b). Pearsall (2000) provides a number of different guidelines for sampling strategies that target either individual excavated contexts such as house floors, or sub- sampling multiple contexts from stratigraphic profiles. Often project research questions dictate the selection of one of these over the other, but an excellent example of integrating both these techniques was published by Coil (2003) in a study targeting changes in agricultural practices and ecology as part of the Kahikinui Archaeological Project in Hawaii. The combination of archaeological and palaeoecological data from individual features, trenches in agricultural landscapes, and sediment cores enabled discussion of landscape use and change within temporal and geographic scales.

Where studies focus on site function, sampling tends to target individual features or contexts of interest within a defined boundary. In the Pacific, Horrocks (and Barber 2005; Horrocks and Bedford 2005, 2010; Horrocks, Smith, Nichol, Shane and Jackman 2008; Horrocks, Smith, Nichol and Wallace 2008; Horrocks and Wosniak 2008; Horrocks et al. 2004, 2012) has sampled a range of agricultural features such as ditches, stone walls and terraces for preserved starch, as well as the surrounding landscape. Balme and Beck (2002) sampled residues on artefacts and sediments to discuss inter-site activity areas within a rock shelter at Petzeks Cave in NSW, Australia. Similarly, sampling of sediments from prehistoric features has been integrated into archaeological projects carried out in Asia and the New World, particularly in locations where macrobotanical preservation is low. This analysis has led to the identification of functional variation within and between sites (Laurence 2013; Messner, Dickau and Harbison 2008), which has been extrapolated further to discus mobility, and the nature and chronology of plant domestication (Barton 2005; Holst, Moreno and Piperno 2007; Messner, Dickau and Harbison 2008; Perry et al. 2007). These examples demonstrate how well-dated and carefully sampled individual contexts can provide information upon site function.

Stratigraphic column sampling, often within test pits or excavation trenches, provides a means of establishing plant use over time. The technique is borrowed from palaeoenvironmental

45 research (Pearsall 2000), and involves evenly spaced or continuous samples taken from the exposed section of excavation units (Coil 2003; Lentfer and Therin 2006; Therin et al.1999; Torrence 2006b). Sampling generally proceeds from the base of the freshly scraped section and works towards the top, with samples carefully placed into labelled zip lock bags using cleaned sampling tools to avoid cross-contamination (Lentfer and Therin 2006:153). Others prefer to sample in the laboratory environment and use box monoliths to collect sediments in situ (Reitz and Shackley 2012). Sampling using paleoenvironmental coring techniques is also common where information is sought upon background environmental and landscape change (Coil 2003; Horrocks et al. 2011; Therin et al.1999) and is validated by studies of modern environmental variation within surface samples (Lentfer et al. 2002; Lenfter and Therin 2006).

Sample sizes often vary as it is difficult to gauge the density of starch preserved within sediments unless a pilot study has been conducted (Torrence 2006b). Most researchers recommend that more sediment is taken than the standard 1-5g that will usually be processed in each batch, with standard sample sizes of up to 100g taken from each context or stratigraphic level (Fullagar et al. 1998:51-2). This enables replication of extraction and identification processes to confirm results, as well as multiple samples to be taken for the extraction of other organic material aside from starch such as phytoliths and pollen.

Once these samples have been collected in the field, they are sub-sampled for starch extraction in the laboratory using one of a number of methodologies. Steps generally involve sample preparation, disaggregation and deflocculation to break up sediments, the removal of unwanted particles, slide mounting and viewing; however, there are a number of ways in which each of these steps can be carried out (Coil et al. 2003; Torrence 2006b). Experimental studies have demonstrated that many chemicals used during standard paleoenvironmental laboratory protocols can damage starch, and therefore alternatives must be sought (Crowther 2009; Torrence 2006b). Crowther (2009) conducted a range of experiments to test a number of chemicals and processing techniques that had been used previously to extract starch from sediments and charred residues. These experiments showed that nitric acid, hydrogen peroxide and heavy liquid in the form of sodium polytungstate (Na6 (H2W12O40)(SPT) had some damaging effects on starch granules. Instead, Crowther (2009:82-83) recommended a protocol using simple chemical disaggregation using weak sodium hydroxide in conjunction with mild agitation and limited sonication within an ultrasonic bath.

Similarly, Torrence and Therin (in Torrence 2006b) tested the effects of Calgon, caesium chloride (CsCl), and sodium polytungstate— two chemicals used for heavy liquid flotation on native starch granules. Samples were monitored at regular intervals up to one week by sub-sampling. The results of these experiments contrasted with Crowther’s (2009) in that sodium hydroxide did not have any corrosive effect on granules, nor did this chemical affect

46 starch quantities when sub-samples were compared over time (Torrence and Therin, in Torrence 2006b:156-7). Caesium chloride was found to have a deleterious effect, but this peaked after five hours of exposure and starch quantities subsequently stabilised. Overall, the 5% dilution of Calgon was found to be the most damaging chemical, with only 59% of starch remaining after five hours of exposure. Torrence and Therin (in Torrence 2006b:157) concluded that other means of deflocculation be explored in the future. The difference between these results for sodium polytungstate may be explained by another study on taphonomy in the laboratory by

Korstanje (2003), where Zinc Iodide (ZnI2) was assessed as another potential means of heavy liquid separation. In this experiment, the use of the chemical was constant, while other variables such as humidity and temperature varied. Korstanje (2003:116) concluded that Zinc Iodide could only be considered destructive to starch when combined with humidity and heat during slide preparation and scanning.

Heat is a variable that has been debated by starch analysts with regard to starch extraction protocols. As discussed earlier, starch gelatinises or melts when exposed to heat and moisture, and so these environments are mostly avoided in starch extraction protocols. However, some protocols suggest that sediments are dried prior to processing to ensure that weights of sediment remain consistent within samples, and also to reduce excessive moisture that can affect the concentration of chemicals (Lentfer and Therin 2006:160). It is argued that heat should be kept at a maximum of 40ºC, when a longer period of exposure is required. This temperature limit also applies to slide drying, where it is recommended that slides are either dried slowly at very low temperatures on a hotplate (Lentfer, pers comm. 2008), or covered at room temperature (Coil et al. 2003; Kortanje 2003).

Variation within processing protocols also reflects the type and contents of the sediments that are being processed. Some chemicals such as hydrogen peroxide are useful oxidising chemicals to disaggregate sediments high in clay or organic content, or charred residues (Crowther 2009; Lentfer and Therin 2006). Calgon is also often used as a deflocculant to separate clay particles. Where these are not required, many protocols simply use heavy liquid flotation and sieving to separate starch from sediments (Atchison and Fullagar 1998; Barton et al. 1998; Fullagar et al. 1998; Messner 2011; Therin et al. 1999; Therin 1994). Many researchers are also quick to point out that techniques are often only in early stages of development and so protocols are only broad outlines of principles rather than definitive methodologies (Fullagar et al. 1998; Korstanje 2003; Therin et al. 1999). Protocols for starch extraction must constantly be updated in light of new data about starch modification that is produced both in the food sciences and archaeobotany. Microwave extraction is a new technique that is currently being tested, especially where fast results are required (Parr 2002, 2006), and involves the pressurization and digestion of organic material within the microwave before sieving to capture microfossils. 47

Chapter 4 Reviewing Parenchyma

The analysis of archaeological parenchyma (or charred vegetative storage tissues) has been under-researched within archaeobotany due to a prevailing view that these remains do not preserve in tropical conditions. Where these remains are preserved in archaeological contexts, they can be extracted through a combination of excavation and flotation practices. When analysed, these data can then provide information upon the diet and subsistence practices of prehistoric cultures that utilise root and tuber crops, such as those in the Pacific. Additionally, the presence of charred parenchyma can be used to infer inter- and intra-site function involving food processing and cooking areas, and also the timing and geographic range of crop dispersal and population migration. Hather (2000) in his seminal guide suggests compiling a comparative collection of relevant domestic and wild species, in the form of both histological thin sections and experimentally charred samples that should be analysed using Scanning Electron Microscopy (SEM).

Fresh and charred parenchyma morphology Few archaeobotanical studies have sought to establish the range of morphological characteristics of vegetative storage parenchyma that are distinctive at various taxonomic levels. Most of what is currently known has been sourced from biology and agricultural sciences. Histological thin sectioning and Scanning Electron Microscopy (SEM) have enabled gross morphology and anatomical characteristics of plant roots and tubers to be studied in detail, and have been replicated by archaeobotanists. Basic anatomical characteristics can be divided into those observed in root-derived tissues and those from stem-derived tissues (Hather 2000; Lebot 2009). A root is the underground portion of the main axis of the plant or branches of the axis, while a stem is the portion of the main axis or branch that is leaf and flower-bearing (Pearsall 2010:153). When stems occur underground, such as corms and stem tubers, they retain many of the anatomical leaf-bearing features including buds and internodes (2010:153; Pate and Dixon 1982:14-21). In contrast, roots lack these gross morphological features. Primary root tissues within dicotyledons and monocotyledons share many morphological similarities (Barlow 1987; Hather 2000:61). A central stele contains the vascular tissues of the phloem, xylem and the vascular cambium surrounded by a layer of endodermis and pericycle enclosed within the outer cortex and epidermis. Secondary root cellular structure in gymnosperms and dicotyledons displays secondary growth within the secondary vascular tissues of the phloem and xylem originating from the vascular cambium, and a periderm originating from an outer cork cambium (Hather 2000: 61; Pearsall 2010:153). Monocotyledons do not exhibit secondary root growth. Stem tissues contrast with roots, consisting of a much simpler anatomy. An outer epidermis surrounds a region of ground tissue or cortical parenchyma, through which runs vascular tissues and the stele (Hather 2000:49). Within these broad anatomical distinctions, taxa differ

48 morphologically due to the function and growth context of these various tissues. Cell shape, size, wall characteristics, and arrangement within the ground and vascular tissues vary across taxonomic levels, along with the arrangement of vascular tissues and cavities within the stele (Hather 2000; Pearsall 2010:158). Patterns of tertiary growth can also differ, sometimes even at the level of individual plants due to localised environmental conditions.

Archaeobotanists by necessity must take these morphological analyses a step further to compare fresh and unaltered parenchymatous tissues with experimentally charred material that is more likely to resemble the preserved botanical remains found in archaeological contexts. A small number of studies have attempted to replicate archaeological charring conditions through the use of muffle furnaces or experimental hearths (Hather 1993, 1994a, 1994b, 2000; Mason et al. 1994; Oliveira 2008, 2012; Paz 2001; Pryor et al. 2013). Samples of vegetative storage parenchyma from species that had geographic ranges pertinent to the locations of archaeological projects were selected, charred under various conditions and then observed using combinations of light microscopy and SEM. Many distinctive changes in morphology were noted; particularly where samples were charred from fresh state and so still had high moisture content, as opposed to being dried prior to exposure to heat (Hather 1994a, 1994b, 2000; Paz 2001).

Alterations of vegetative storage tissues occur during charring or desiccation, but vary according to charring conditions. Variables such as temperature, period of exposure to heat and oxidising conditions (such as immersion in substrate) can create further modification of tissue morphology (Hather 1991:663; Hather 1993; Pearsall 2010). Features such as the creation of tension fractures and vesicles, the deterioration and subsequent transformation of the phloem into cavities or solid carbon, and modification of cell characteristics such as wall thickening and compression can alter the original morphology of roots and tubers (Hather 1991:3-8, Paz 2001:88-105). Tension fractures are caused by the ripping apart of tissues along lines of weakness, while vesicles are created through the dissipation of steam within the tissue (Hather 1994a:54). Lignified tissues such as the xylem are generally preserved during charring, but the living tissues within the ground tissue and the phloem are either altered or completely breakdown. The water content of tissues prior to charring can significantly increase the extent to which these tissues are altered or damaged (Hather 1993:3). It is important that the nature and scope of these changes is understood when attempting to identify unknown botanical remains extracted from archaeological or palaeoenvironmental contexts.

Taphonomic factors affecting macrobotanical preservation Unlike the analysis of starch granules within archaeological and palaeoenvironmental contexts, the taphonomy of vegetative storage parenchyma is poorly known. This is a direct result of the lack of archaeobotanical projects targeting the charred remains of these relatively soft tissues under the presumption that preservation will be low or non-existent. Conditions for preservation

49 of parenchyma are primarily limited to those that either desiccate, water-log or char these botanical remains (Hather 1991, 1992, 1994, 2000). Another less common form of preservation includes mineralised or semi-mineralised remains that are either impregnated or coated by semi- crystalline minerals that protect the soft tissues from fast decay (Paz 2001:40). While wood charcoal and seeds may change little on preservation (unless compressed), vegetative storage tissues tend to undergo a number of physical transformations (Hather 1991:662). This is because roots and tubers have a higher proportion of parenchyma tissues and a smaller portion of cells with lignified walls than stem wood (Pearsall 2010:161). Experimental charring has emphasised the nature of some of these changes which include expansion or shrinkage of tissues, the deterioration or loss of more fragile regions, and fragmentation (Hather 2000:74, 1991:662-3).

The paucity of root and tuber macro-remains in archaeological sites is often directly related to how these resources are processed and used. As these are generally sources of food the only part of the organ that often remains is the peel (Allen 1983; Kahn and Ragone 2013; Pearsall 2010:154). Some root peelings are tough and fibrous but others have periderms that decay relatively quickly and are very fragile if charred. Other aspects of the biology of roots and tubers also make any preservation unlikely. For example, if these organs are dropped or discarded by humans they are often scavenged by animals or are susceptible to agents of decay due to the high calorific value and moisture content (Holden et al. 1995:777). High moisture content also makes these organs prone to distortion and fragmentation if exposed to heat, especially once charred (Holden et al. 1995:777; Pearsall 2010:157). Those fragments or whole organs of vegetative storage parenchyma that do preserve are most likely roots and tubers that have been discarded into the fire as spoiled goods or accidentally charred during roasting (2010:157). Ethnoarchaeological observation in the Philippines has confirmed these as possible cultural taphonomic processes that enable charred parenchyma to enter the archaeological record (Paz 2001:80). Charcoal is formed in the reducing conditions within the ash in the base of a fire or hearth, rather than the oxidising conditions of the open flame which eventually reduces tissue to ash. Some small dense fragments of tissue can fall through the structure of the fire into the ash and transform into preserved charcoal (Hather 1991:663).

Hather warns that quantification of taxa within preserved parenchyma is unlikely to result in meaningful interpretation because the cultural and taphonomic processes affecting the conditions of preservation are largely unknown (Hather 2000:74). However, it is possible to utilise some of the current data about macrobotanical preservation and post-depositional processes derived from studies of wood charcoal or anthracology. Thery-Parisot and others (2010) describe the range of ‘filters’ that botanical remains will pass through before being incorporated into palaeoenvironmental or archaeobotanical reconstructions. These include societal filters such as selection, hearth maintenance and storage; combustion filters such as anatomical changes and differential fragmentation; depositional and post-depositional filters 50 include anthropogenic factors, mechanical factors and diagenesis; and archaeological or anthrocological filters including sampling, identification and quantification (Thery- Parisot et al. 2010:143). Post-depositional filters can be broken down into specific cultural processes such as trampling, re-working and sweeping, and natural taphonomic agents such as bioturbation, atmospheric factors, mechanical constraints that cause pressure or friction in the sediments and can induce chemical alteration, water and the pH of soils (Thery-Parisot et al. 2010:147-150). Where these process act homogenously within a particular context, they will not affect the palaeoecological signature, but this is rare and differential preservation can often be observed within these records.

These post-depositional processes can lead to vertical and horizontal migration of remains as well as fragmentation and disappearance (Paz 2001, 2005; Oliveira 2008; Hather 1994, 1995). Nelson (1992) and Greenlee (1992) assessed the nature of downward displacement of macrobotanical remains within shell midden contexts. Each argues that the ratio of shell to soil matrix will affect the porosity of the midden, and thus also the amount of movement that is possible (Greenlee 1992:262; Nelson 1992:254). As a general rule, particles smaller than 2mm are most susceptible to movement through mechanical processes, as well as bioturbation by roots, earthworms and burrowing animals (Nelson 1992:254; Thery-Parisot et al. 2010:147). Paz (2001) also explores the role that soil matrix can play in the preservation and movement of macrobotanical remains. He argues that the compaction of clay soils inhibit matrix mixing except where bioturbation has occurred, but agrees with Nelson (1992) and Greenlee (1992) that the porosity of shell midden and also sandy deposits allow greater degrees of turbation and downward displacement (Paz 2001:40).

Questions of context security can also be addressed through assessment of the forms and taxa of preserved macrobotanical remains within a given strata. Purely charred remains can be considered reasonably representative of an in situ deposit, especially where upper layers do not contain any untransformed or intrusive remains that have not undergone charring, desiccation, water-logging or mineralisation (Paz 2001:262). Where downward percolation has occurred, the distribution of seeds and small charcoal should be indicative of this movement, with greater densities of remains in lower levels of particular deposits or facies within shell middens (Nelson 1992:243). Post-depositional homogenisation by chemical processes such as prolonged submersion in brackish groundwater can also occur and mask variability in separate and disparate depositional events (Nelson 1992:251). These studies argue that by assessing the context, sedimentary matrix, and nature of botanical remains, some natural and cultural post- depositional taphonomic processes can be recognised and incorporated into the interpretation of the archaeobotanical record.

Macrobotanical preservation within tropical climates like those in the Pacific is generally thought to be poor. However, a number of archaeobotanical projects in the Asia- Pacific region have proven that these conditions do not necessarily inhibit long term preservation of soft tissues (Allen 1983; Barton and Paz 2007; Hather 1991, 1992, 1994a, 1994b, 1995; Paz 2001, 2005). Desiccated remains are rarely found unless in caves or rockshelters. Water-logged remains are a more common form of soft vegetative tissue recovered in the region (Hather 1992:71). Anaerobic brackish lagoon settings have preserved large assemblages of botanical remains within modern beach deposits formed by clay erosion. For example nuts and other large seeds were preserved within these conditions at the site of Talepakemalai on Eloaua Island in the Mussau Island group (Hather 1992; Kirch 1989). Charred remains of roots, tubers, wood, seeds and nuts form the majority of preserved plant tissue found in archaeological and palaeoenvironmental contexts. Within these remains, fruits, roots and tubers are less common, but have been extracted and identified on a number of occasions within rock shelter deposits (Hather and Kirch 1991; Oliveira 2008, 2012; Paz 2001), shell middens, house floors (Coil and Kirch 2005; Kahn and Ragone 2013; Yen 1974) and dry sandy open sites (Hather 1994).

Collection and sampling of parenchyma Preserved vegetative storage parenchyma can be extracted from archaeological contexts using established archaeobotanical methods for collection of macrobotanical remains. These usually involve flotation, or wet or dry-sieving of excavated material, alongside collection of material in situ (Hather 1994a; Pearsall 2010; Thompson 1994:14). Sieving involves the manual agitation of material to loosen sedimentary aggregates and allow smaller particles to fall through the various sized meshes, retaining a portion that contains botanical material (Hageman and Goldstein 2009:2846).This separation is more effective and less biased than collection of botanical material by eye during excavation. Dry-sieving is not always effective when processing clay or silty sedimentary matrices, and so wet-sieving or flotation is more commonly used in these circumstances. Wet-sieving utilises water to further breakdown tough aggregates or sediments that have high clay content. Limiting factors affecting the effectiveness of dry or wet-sieving techniques are the size of the mesh used which can bias the types of remains collected, and the amount of force used to push material through the mesh can also damage or fragment preserved botanical remains (Pearsall 2010:13).

Water flotation utilises differences in density of organic and inorganic material to separate organic material such as charred botanical remains from the soil matrix. When carried out on a large scale, this technique can separate much higher quantities and range of botanical material than sieving alone (Pearsall 2010). Mechanical bulk flotation devices can be employed to process large amounts of sediment where water is freely available or where equipment is available for water recycling (Hageman and Goldstein 2009; Pearsall 2010); however, these 52 machines can increase the risk of charcoal fragmentation. These systems use flowing water to break up sediments and release any charred botanical remains that have a specific gravity or are less dense than water to float to the surface. The remains are caught in sieves with varying mesh diameters to create flot fractions of different sizes (Pearsall 2010:50-52). Mechanical systems are designed to limit the need for extra personnel to operate them, but must be overseen for maintenance and to collect samples (Hageman and Goldstein 2009:2847-8). Bucket flotation is often used where equipment and access to water is limited (Fairburn 2005b; Mason et al.1994; Oliveira 2008; Paz 2001). These tub or bucket systems involve manual agitation of sediments, with muslin cloth pegged into a bucket replacing sieves to catch botanical remains that are decanted from the surface of the diluted sediment. Heavy residue remains in the bucket, which is then either discarded or wet-sieved, to collect any remaining botanical or artefactual material. Comparisons of the flotation techniques have demonstrated the effectiveness of the mechanical systems (40-100%) over the bucket systems (6-100%) in terms of recovery rates (Wright 2005).

It has been argued by some archaeobotanists that dry-sieving is the most appropriate technique for the separation of vegetative storage parenchyma due to the fragility of these soft tissues when preserved in charred, desiccated or mineralised forms (Hather 2000:74). Other studies have demonstrated that the utilisation of both sieving and flotation can result in the recovery of increased quantities and also greater taxonomic diversity (Fairburn 2005b; Hageman and Goldstein 2009). Both techniques have advantages in the separation of particular types of botanical remains and specific taxa. Flotation enables the collection of material with low specific gravities such as many seeds, while sieving can facilitate the collection of endocarp and wood that is often too dense to be recovered through flotation. A combined approach is essential when consideration is given to the fact that that different environmental conditions during carbonisation can cause some botanical remains to vary in density, and thus affect the odds of recovery through flotation (Wright 2005:24).

The selection of a sampling strategy can also influence the recovery of botanical remains using techniques highlighted here. It is most often impractical to process all excavated material from within large-scale projects for all size-classes of preserved botanical remains, and an appropriate sampling regime must be chosen that enables the particular research questions to be answered. Lennstrom and Hastorf (1995) caution against taking the ‘feature bias’ approach that targets only features or areas of concentrated carbon, as it is just as relevant to discover areas that do not contain botanical material. The most common macrobotanical sampling techniques are the ‘blanket sampling’ strategy, whereby a sample is taken from every excavated context and feature (Fairburn 2005b:10; Pearsall 2010:66-68). These include ‘scatter’ or composite sampling where small amounts of matrix are gathered throughout a context and combined in the same collection bag, where samples derive from a column or sequence of deposits, and point bulk sampling of precisely located areas within a context (Pearsall 2010; 53

Wright 2005:20). Comparison of these techniques indicate that composite samples tend to recover higher quantities and diversity of charred material, but had a smaller range of variability within the density of material compared to point bulk sampling. Pearsall (2001:71) argues that this indicates that scatter samples captures a greater range of actions and places within an occupation level, but may also be a biased approach to sampling.

Sampling strategies involve the collection of a small amount of material considered representative of a particular context, but the way and form in which the sample is measured can also vary, and influence archaeobotanical interpretation. Sample size is usually measured by weight or volume, although these can both be influenced by the amount of moisture within the sedimentary matrix (Wright 2005:20). Even very small amounts of water can increase the weight of soil, for example 1ml of water at 4ºC weighs 1g. Volume is often measured using a calibrated bucket, but wet matrix can be harder to pack into these buckets and thus more difficult to accurately measure. Wright (2005) demonstrated this difference in volume by also comparing the types of sediment being measured when wet and dry or partially dried. Clay sediments had an average increase in volume of 25%, while silty sediment only increased 9% and sand was almost exactly the same, with an increase of only 1%. These results indicate that consistency is required to ensure that sample sizes can be considered representative and are comparable (Lennstrom and Hastorf 1995:705). Despite this, repeated wetting and drying of samples is not recommended, as this can weaken the cellular structure of charred remains and increase rates of fragmentation (Greenlee 1992; Hather 2000; Paz 2005).

Parenchyma identification Researchers, both in the Pacific and elsewhere, targeting the extraction and identification of vegetative storage tissues have emphasised the need to define the criteria upon which taxonomic identification is based. At the most basic level, charred vegetative storage parenchymatous tissues can be distinguished from wood charcoal based on several gross morphological and anatomical characteristics (Hather 1991:673; 1993:3, 2000; Pearsall 2010). These include:

 Fragments are often rounded in shape, unless recently fractured, as corners and protruding tissues tend to be removed cell by cell.  Cells are mostly rounded in shape and more or less isodiametric in dimension, with very few highly elongated cells resembling vessels or tracheids in wood.  The texture of parenchymatous tissues is usually dull but some areas will be dense and reflective where vascular tissues or schlerenchyma has formed solid carbon.  Charred vegetative parenchymatous organs often contain regular or irregular cavities, caused by the evaporation of moisture, that are identifiable under low power microscopy.

Beyond these basic characteristics of charred vegetative parenchymatous tissues, the preservation and size of fragments determine the level of taxonomic identification that can be achieved. Archaeobotanists are currently divided over the diagnostic value of morphological and anatomical characteristics, and how to define confidence in classifications based on these attributes. Most agree that by studying the overall form of root and tuber material, external characteristics, anatomical structure, and alteration of tissues through charring process, it is often possible to identify archaeological material (Paz 2001; Pearsall 2010), but caution against attempting precise identification on poorly preserved remains that lack features of diagnostic value. Hather (2000:72-73) argues that for a character to be diagnostic within the scope of a comparative collection, it should satisfy four conditions. Firstly, it should be common enough to be found in all recovered fragments that have this characteristic. Secondly, it should be easily observable and thus have survived preservation and post-depositional processes. Thirdly, it should be a stable character that does not vary infraspecifically. Finally, the character should vary between species or higher level taxonomic groups. High confidence identifications are not usually based on single diagnostic characters; rather remains are classified using a range of different attributes.

Pioneering research by Hather (1991, 1993, 2000) established the overall diagnostic value of the morphological and anatomical components of roots and tubers through analysing the rate of preservation, frequency of occurrence within an organ, stability and range of variation of each. In general he argues that characters of the gross morphology and surface features are rarely found on preserved fragments of parenchymatous tissues from roots and tuber, unless these organs are small before charring (Hather 2000:73). However, when these are present these are generally of high diagnostic value. Parenchyma and sclerenchyma cells usually preserve relatively well, but the range of variation between these is often observed to be quite low when comparing different taxonomic groups. Vascular tissues also preserve well except within the secondary growth of roots, but in contrast to ground tissue, mostly varies between species or higher taxonomic groups.

Paz (2001, 2005) takes these findings further by establishing the range of variation within the periderm, examples of which have been recovered from several locations around the Asia-Pacific region (as mentioned earlier), including within his own research. Importantly, Paz also describes the specific criteria by which remains can be identified to plant taxon, specific organ or as vegetative storage parenchyma (Paz 2001:82-85). For example, remains can be identified to a particular taxonomic group if the fragments fit all or most of the features of a reference species such as cell size, shape, the arrangement of vascular tissues, and crystal or starch grain morphology (Paz 2001:82). Without preserved cellular structure with observable shapes, walls, and vascular tissues, remains cannot be identifiable to species or any higher taxonomic grouping with high confidence. Instead, prefixes such as ‘cf.’ or the suffix ‘type’ can 55 be used to indicate moderate confidence classifications. When these criteria are not met, these can often still be identified morphologically and artefactually using characters of charring as root or stem tubers (Paz 2001:83). Aside from morphological and anatomical characters, other criteria include matching images, illustrations, local or regional flora citations, taxonomic details of the species, and the geographic distribution of taxa (Paz 2001:71). These criteria established by Paz for the identification of parenchyma have not been universally adopted by other researchers, but Oliveira (2008, 2012) has used the same methodology in his research in East Timor. These criteria were successful in aiding the identification of a number of economic and supplementary crops within these locations.

Chapter 5 Comparative Collection and Morphometric Studies of Pacific Cultigens

Chapters 3 and 4 of this thesis highlighted the issues and potential applications of starch and parenchyma within archaeobotany in the Pacific. The biological features of these botanical remains were outlined, which influence both taxonomic classification and also long term preservation. Preservation of micro and macrobotanical remains particularly in tropical climates was discussed, along with other taphonomic factors such as bioturbation and the effects of cultural processes such as charring and boiling. A number of techniques for the extraction of starch and parenchyma from sediments were also reviewed. These reviews influenced the decision to establish a comprehensive reference collection for this study, which incorporated a detailed morphometric study of both economic and supplementary species from the Kingdom of Tonga.

Species selection Ethnobotanical and historical resources were used as guides for the inclusion of species in the modern comparative collection for this study. The list of crops recorded by early explorers and missionaries such as Cook (1785), La Perouse (1788), La Billardiere (1793), Wilson (1799), Mariner (in Martin 1991), Waldegrave (1873), Gifford (1929), Beaglehole and Beaglehole (1941), and others on Tongatapu, is vital to the development of a comprehensive understanding of late prehistoric and contact period agriculture in Tonga. Ethnobotanists have attempted to assess the origins and dispersal of these plants in the Pacific for many years (Whistler 2009, Yen 1974) and lists of endemic, native, and Polynesian or European introduced species within Tonga are constantly evolving. An inclusive approach was taken in this study, and so sourced many economic species that may have reached the Tongan archipelago. Other researchers have established the value of using ethnographic data as a cautious analogy for the past (David and Kramer 2001; Wylie 1985), and this information was used to develop an ethnographic baseline for the development of a comprehensive comparative collection in this study.

Historical and contemporary sources indicate that prehistoric horticulture within the Tongan archipelago was dominated in late prehistory by a small number of starch-rich crops for the provision of crucial carbohydrates. The most important of these were a range of yams (Dioscorea spp.), including D. nummularia, D. esculenta, D. alata, and D. bulbifera. Dryland production of aroids such as Alocasia macrorrhiza, and Colocasia esculenta were seasonally alternated with the production of yams within plantations, as was the planting of sweet potato (Ipomoea batatas) in mounds. Arboricultural production of bananas (Musa spp.), breadfruit (Artocarpus altilis) and the Tahitian chestnut (Inocarpus fagifer) was also an essential agricultural component. Supplementary to these core species, was a range of alternative plants

57 that were either cultivated or utilised such as the Ambarella (Spondias dulcis), the elephant foot yam (Amorphophallus campanulatus) and the Polynesian arrowroot (Tacca leontopetaloides). These included a number of species that were also part of this transported landscape from Wallacea and Southeast Asia, but others were native or endemic species that produced roots, tubers, fruits and seeds that were either edible through processing, or used for other cultural purposes (such as kava or Piper methysticum, and the fish-poison tree- Barringtonia asiatica). Plant products fulfilled a number of vital roles within Polynesian society, and so the inclusion of a range of species within the comparative collection for this study will shed light on the antiquity of some of these practices with Tongan prehistory.

Field collection The major portion of the comparative collection used in this study was collected from the island of Taveuni in Fiji. Approximately 25 species were collected here, with another 15 species collected in Tonga and Palau. Each vegetative storage organ of the plant was sampled. These species included a range of both economic and supplementary plants. The comparative collection built on an earlier range of economic species that had been compiled for the author’s research in the Marquesas (Ussher 2009). Most islands in Polynesia share the core suite of crops thought to have either been brought as part of the initial Lapita package (Kirch 1984; Green 1979) or through later migration and interaction, with some environmental restrictions upon their production capacities. It is therefore relevant to include most of these species from Palau, Fiji and the Marquesas, in addition to the Tongan specimens, within the comparative collection for this study.

The major families collected included a range of aroids (Araceae), members of the Dioscoraceae and Convolvulaceae family, and Moraceae. A survey of Taveuni was carried out under the guidance of Botanist Bee Gun from the Crisp Lab at the Australian National University. The help of Botanist and Ecologist, Dr Ann Kitalong, was enlisted in Palau to survey areas in the Rock Islands and Babeldoab for cultivars. Local names of species were recorded and used to source plants in communities and plantations. Voucher specimens were collected for each species included in the reference collection, and are stored either at the University of Auckland, or the National Herbarium in Canberra. These were air dried quickly using dry heat in the field, stored within labelled folds of newspaper and compressed using a wooden plant press (Prebble, pers comm. 2011). The dry heat prevented any mould from developing upon the specimens from the humid tropical climate.

The organs sampled for starch were stored for transport within alcohol in order to preserve the original structure (integrity) of the sample, and prevent decay. The samples were then each labelled and sealed with masking tape to prevent leakage and evaporation during transportation and storage until laboratory processing commenced back at the Australian

National University (ANU). A sample collection sheet was completed for each species, including details about the geographic location of each plant, a plant morphological description, and a more specific description of each of the organs sampled using current botanical terminology. In addition to these specimens collected by the author, a selection of other economic cultivars was included from a collection housed at the ANU by Douglas Yen. Samples had been desiccated and powderised before being stored. The desiccated state of these specimens meant that these could only be included in the starch component of the comparative collection. Likewise, a number of charred specimens stored at the ANU by Jon Hather were included in the parenchyma component of the comparative collection.

Laboratory processing of samples Each of the samples collected in the field was first removed from their original containers and put into a new sterilised vial with 70% Ethanol and 30% De-ionised water for storage until further processing began. Each sample was then divided into four sub-samples to allow for starch extraction, histological thin sectioning and experimental charring for parenchyma from dried and fresh states.

Figure 5.1 Flowchart showing methodology for the imaging and recording of starch and parenchyma within the reference collection

Starch processing The starch samples were processed using a methodology established by Ussher (2009, 2012) in creating a comparative collection for the Marquesan archipelago, and including methods suggested in Field (2006). Two methods were used to extract starches from the parenchymatous tissues of the vegetative organs sampled. The first of these were to simply cut a fresh section from the organ and press that section onto the slide (Gott in Field 2006). The second was to gently crush the sample in a mortar and pestle with some distilled water, and then put this 59 extracted residue onto a slide using a pipette (Lentfer, pers.com). The slide was then covered with a Petri dish until the residue had dried. Glycerol was used as a permanent mountant, due to the reflective and viscous properties of this compound which allows starch granules to be rolled and viewed during light microscopy (Field 2006:112). Finally, each sample was covered with a cover-slip and sealed with nail polish.

Histology A histological thin section was made from each plant species and organ collected, to demonstrate parenchymatous cell arrangement, structure and contents. The samples collected were removed from the 70% Ethanol, and a small 1cm cubed fragment was taken from each sample and placed into a mixture of 95% (70% dilution) alcohol, 1% glacial acetic acid, and 4% formaldehyde. This fixative formula was recommended by Hather (2000:78) and is known as FAA (Formalin-Acetic-Alcohol) (Miksche and Berlyn 1976:30). This fluid is stable, has a good hardening action and material can be stored in it for years. The samples were left in the FAA mix fixative for approximately two weeks, by which time the organs had soaked up the mixture and partially solidified/preserved as dead plant tissue. This solution enabled greater precision during microtomy to cut thin sections.

Thin sections were made by Anne Prins in the Histology Laboratory at the John Curtin Medical Research Centre. Several methods for processing samples prior to microtomy were tested. A white potato (Solanum tuberosum) was also preserved in the fixative formula, and then used to experiment with microtomy and staining. The Standard Bouins Cycle technique was applied to prepare the samples after they were placed into small histological baskets. This technique involves several stages of soaking samples in varying percentages of ethanol warmed to 40˚C, starting with 70% and finishing in 100%, then baths of chloroform at 40˚C and paraffin wax at 60˚C. These were then placed in the microtome and cut into 5, 8, 12, 15 and 20µm thin sections. Experimentation with the white potato indicated that the most appropriate section size during microtomy was 15µm, as thinner sections tended to rip and thicker sections were difficult to mount onto the slides. The paraffin embedded thin sections were then mounted on heated slides and placed in xylol to dissolve the wax, and finally washed with 100%, 90% and 70% ethanol, and tap water prior to staining.

As part of the staining process the sections were treated with 3% glacial acetic acid for three minutes, and then washed well with running tap water. The slides were then immersed in Alcian Blue (1%) stain with a pH of 2.5 for 30 minutes, washed in running tap water again, and finally immersed in a Safranin (0.02%) counter-stain for five minutes and then dehydrated. After staining, the sections were blotted, allowed to air dry and cover-slipped with Leica Micromount used as a permanent mountant.

The experimental thin sections were observed using light microscopy at x100 and x200 magnification. The cell walls were in good condition, and many starch granules were still visible and displayed birefringence in some samples. The fact that starch was preserved in these thin sections was surprising, as starch generally gelatinises at between 30-60˚C (Barton and Torrence 2006). Here the starch was not only still in native condition, but also abundant within the cell structures. It was therefore decided that the Paraffin embedded preparation was the most appropriate for the remainder of the samples within the reference collection. The samples were prepared in the same way as the experimental white potato and then analysed using light microscopy in the Microscopy Laboratory in the Department of Archaeology and Natural History at the ANU.

Experimental charring To understand the morphological changes that occur during charring, plant materials within the reference collection were experimentally charred in a muffle furnace. These samples were then compared with the parenchyma cell organisation and structures observed in the thin sections. Recommended temperatures and length of time for charring vary according to the type of plant tissue, i.e. woody or non-woody tissue. Orvis and others (2005) suggest charring woody stem material, soft-leaf tissue, needle-leaf tissue and monocot tissue at 550˚C for only eight to nine minutes. Boardman and Jones (1990) tested a range of temperatures, times and atmospheric conditions (oxidised or reduced). It was discovered that most plant components (granules, glumes, rachis and straw) will carbonise after 1-2 hours at 300-250˚C, with greater distortion at higher temperatures even within shorter time periods (1990:5-9). Similarly, Wright (2003) experimented with charring achenes, kernels, seeds and rind segments of a variety of species. Variables tested included thermal atmosphere, temperature, duration and moisture content. Overall, some generalities could be made regarding the carbonisation process including that the higher the temperature, and longer the exposure, the more likely the specimen will ash rather than carbonise, and that moist specimens generally survived oxidation better than dry samples. Finally, a reducing atmosphere was more conducive to preservation by carbonisation than oxidation (2003:582). The results of these previous studies were used to determine the techniques used within the current research.

Two small 1cm cubed samples were cut from the remaining plant material for each sample, and wrapped in tin foil. One sub-sample of each sample was placed into large tin trays, and then put into an incubator at 60˚C for 48 hours to dry. The second sample was placed into a pre-heated muffle furnace at 300˚C for 2 hours (Hather 2000:85), or until the samples had turned to charcoal (Boardman and Jones 1990:3). They were then removed from the furnace and cooled. Once dried, the remaining samples were also charred in the furnace. All samples were then fractured to reveal a flat plane that could be viewed using Reflected light microscopy and Scanning Electron Microscopy (SEM) (Hather 2000:76). 61

Recording

Light microscopy

Starch Light microscopy was used to view and record attributes of both starch residues and also parenchymatous plant material in the comparative collection. A Leica DM6000 Compound Transmitted Light Microscope was utilised to analyse the modern starch samples and record 18 attributes of 50 granules from each slides. Images of each grain were taken, and these were then analysed using ImageJ software to measure specific attributes. These attributes included maximum length, width, length of the extinction cross, distance between the hilum and end of the grain, maximum distance between the arms of the cross, maximum angle of the cross, hilum location, 3D shape, and presence of lamellae, a small or large vacuole, an equatorial groove, compressed or discoid structure, multiple facets, flat facets, hilum fissures, raphides (calcium oxalate crystals), and the style of extinction cross.

The inclusion of metric variables has been recommended by a number of previous morphological studies including that by Torrence and others (2004), and more recently by Wilson and colleagues (2010) in their study using image analysis to determine morphological variety within native reference starch. Metric variables enable much greater differentiation between taxa, and reduce observer bias during classification. Granules were viewed in both brightfield and cross-polarised light, so that attributes of the extinction cross could be imaged and recorded. The data collected during light microscopy was then entered into an Excel database, with a spreadsheet created for each species and organ sampled.

Parenchyma The histological thin sections were also viewed using light microscopy. Both transverse and longitudinal sections were imaged and attributes of the cell structure and arrangement were recorded. These included descriptions of the boundary tissues such as the epidermis, periderm and cortex, general ground or conjunctive tissues, as well as the vascular tissues. Cell morphology was recorded through attributes such as cell length, width, shape and dimensions, as well as cell contents and the presence of inter-cellular spaces. The arrangement and organisation of vascular bundles were also recorded according to known typologies (Hather 2000). To gain representative samples of tissues, 40 cells from each species and organ, and five vascular arrangements were measured and described. The images of the thin sections were viewed and analysed using ImageJ software.

Scanning Electron Microscopy High resolution, low voltage field emission scanning electron microscopy (HRLVFESEM) was chosen as one method for the analysis and imaging of the charred parenchyma and starch samples. This technology provides topographical contrast of small features that are only

62 minimally coated while charging is substantially decreased as a result of lowered beam energy (Schatten 2008), creating a three-dimensional view of objects. Secondary emission (SE) signaling was chosen as the amount of SE signal collected from each point is roughly proportional to the angle between the viewing direction and the normal surface. This type of image conveys a relatively accurate impression of sample surface topography to the human brain (Pawley 2008), creating a detailed three-dimensional image of important morphological features.

Sample preparation The experimentally charred samples were carefully hand fractured to create a flat plane for viewing cell structure and arrangement, then attached to an SEM stub using carbon tape. All starch samples were processed in the following way. A small amount of crushed specimen was mixed with distilled water onto a small circular glass cover-slip, which was then attached to the stub using nail polish and left to dry. Carbon tape was placed over the edge of the cover-slip and the sides of the SEM stub to create a path for the electrons to escape when the sample was placed in the SEM chamber. All samples were sputter coated with platinum gold at 20 milliamps for 3 minutes to create a relatively even conductive coating for imaging and analysis.

Imaging The Zeiss UltraPlus FESEM at the Centre for Advanced Microscopy at the ANU was then used to image each sample at up to 10,000x magnification. A total of nine samples were able to be placed in the chamber of the SEM at any one time. Each sample was targeted in turn and images taken of the sample at various magnifications, with appropriate scales embedded. Lower magnifications of 200-1000x were used to capture images of the cellular tissues and their arrangements. Higher powered magnification was used to take high-resolution images of cellular structure, cell contents, and modifications that occur during the charring process. Imaging was affected by occasionally insufficient or uneven sputter coating of the sample resulting in the charging of the electron beam which causes images to be blurred in areas. This could not be avoided in some instances as re-coating with platinum gold would result in further unevenness and charging. The resulting images of the starch and charred samples of parenchyma were used to supplement the descriptions of morphology within the comparative collection, and were vital towards enabling identification of unknown archaeological material in Part Two of this thesis..

Morphology of native starch Research within the food sciences has been investigating starch chemistry and physical morphology for many years. The results of these studies have demonstrated that these aspects of starch can vary at different taxonomic levels; however it is clear that many species contain a number of starch morphotypes that are shared with other species (Banks and Greenwood 1975; Crowther 2012; Henry et al. 2009; Nwokocha and Williams 2011; Parr 2002; Reichert 1913). 63

The compilation of a comprehensive reference collection including both economic and wild starch-producing species is essential to identify starch preserved within archaeological contexts. Understanding the characteristics that differentiate between starch morphology at species, genus or family level, is the only means by which starch extracted from archaeological sediments or as residues on tools can be securely identified.

Starch morphology As mentioned previously, a range of starch attributes were analysed for all plant specimens in the comparative collection. These included a combination of metric, nominal and binary attributes. It became clear that each specimen (species or organ) contained a range of different starch morphotypes. Sometimes as many as eight morphotypes were recorded within a single sample. A signature type was usually present in each specimen and was recognised as being the most diagnostic morphotype within the random 50 granules recorded. This may not necessarily be the most commonly represented morphotype. It will be demonstrated here that there is a large degree of morphological overlap between many specimens within the modern comparative collection (see Table 5.3). Nonetheless, the identification of these signature types is essential for establishing an explicit method for the taxonomic classification of unknown archaeological starch granules.

Hilum features The hilum is the point from which a starch granule grows and is thus the core of the granule (Banks and Greenwood 1975). Due to the fact that the hilum is a fixed point on the granule, its location appears to change as the granule is rolled and viewed from different angles. The hilum can therefore be used as a point of orientation when describing the three-dimensional shape of granules. A number of genetic modifications can alter the appearance of the hilum. Hila can often be observed as a small or large vacuole, or as a fissure or crack on granules. In the comparative collection developed for use for this study, a large range of specimens contained starch with a vacuole at the hilum (see Table 5.1, Figure 5.2). Small vacuoles were recorded on starch morphotypes within 18 specimens, and were commonly seen in 11 of these. All recorded starch from Piper methysticum had small vacuoles. Conversely, small vacuoles were rare in most of the Artocarpus and Dioscorea genera, as well as in the seeds of Barringtonia asiatica and Pteridium sp. Large vacuoles were present in a much smaller number of specimens, and were common in M. citrifolia and S. dulcis, but rarely observed in B. asiatica, Cyrtosperma merkusii and Ipomoea polpha. The starch morphotypes of Spondias dulcis all had either a small or large vacuole, indicating that these are diagnostic traits of starch from this species.

Figure 5.2 Diagram showing basic features of starch granule morphology

Fissuring at the hilum was also observed within 18 specimens in the comparative collection. There are a range of different types of fissuring that can occur within starch (ICSN 2011), and a small number of these were present within the collected specimens. The most common type was longitudinal fissuring, where the fissure extends along the long axis of the grain. Nine specimens had granules with longitudinal fissuring, but it was only common in three species, Amorphophallus paeoniifolius, Dioscorea rotundata and P. methysticum. Radial fissuring was also identified. This is defined as fissuring that originates at the hilum and spreads outwards to the margins (ICSN 2011; Reichert 1913), and was commonly recorded within two species, Ipomoea polpha and Piper methysticum. Transverse fissuring extends at a right angle to the long axis of the grain (ICSN 2011), and was observed in a total of five specimens (see Table 5.1). Both I. polpha and Spondias dulcis granules commonly feature this hilum type. Stellate fissuring is another common type recorded within the comparative collection. These star-shaped fissures were common in three species- I. polpha, P. methysticum and S. dulcis, and rare in Artocarpus heterophyllus and Inocarpus fagifer.

Several other types of hilum fissuring were identified within a smaller range of species. Oblique fissures are simple fissures that do not follow any particular axis on the grain (ICSN 2011, Reichert 1913) and were found commonly on granules from Dioscorea rotundata but only very rarely on Solanum tuberosum. Irregular fissures are those that are uneven in geometry and were observed on the starch of three specimens; however, this type was only common in I. polpha and P. methysticum. The presence of a mesial cleft, a large, deep and variably ragged interior crack that runs parallel to the long axis of the grain (ICSN 2011), was only noted in the starch of P. methysticum. Likewise, branching fissures were only observed in S. dulcis. These are fissures that feature subdivisions from the main branch. The final type of fissuring within the comparative collection is actually classed as an ‘indentation’ at the hilum, and was observed frequently in A. paeoniifolius but only rarely in I. fagifer.

Longitudinal Radial Oblique Amorphophallus paeoniifolius Artocarpus heterophyllus Dioscorea rotundata Artocarpus altilis seed Barringtonia asiatica fruit Solanumtuberosum Dioscorea nummularia Cyrtosperma merkusii Dioscorea rotundata Dioscorea alata Indentation Inocarpus fagifer seed Dioscorea bulbifera Amorphophallus paeoniifolius Musa sp. 2 Ipomoea polpha Inocarpus fagifer seed Piper methysticum Piper methysticum Transverse Tacca leontopetaloides Barringtonia asiatica fruit Xanthosoma sagittifolium Stellate Dioscorea rotundata Irregular Artocarpus heterophyllus Ipomoea batatas Cyrtosperma merkusii I.fagifer seed Ipomoea polpha Ipomoea polpha Ipomoea polpha Spondias dulcis Piper methysticum Piper methysticum Branching Mesial cleft Spondias dulcis Spondias dulcis Piper methysticum

Table 5.1 Hilum fissuring of reference species

Lamellae Lamellae are the growth layers that begin forming at the hilum and disperse towards the margins of the granule. These were ‘distinct’ in many large starch morphotypes deriving from the Dioscorea genus including Dioscorea alata, Dioscorea bulbifera, Dioscorea nummularia, Dioscorea pentaphylla, and Dioscorea rotundata. Distinct lamellae were also visible on all recorded starch from Curcuma longa and most starch from Musa sp. 1. ‘Indistinct’ or less prominent lamellae were observed on larger starch morphotypes from the fruit of Artocarpus altilis, Horsfieldia palauensis, Ipomoea batatas, Ipomoea polpha and Musa sp.2 .Smaller starch morphotypes, such as those seen within Alocasia macrorrhiza, Colocasia esculenta and Dioscorea esculenta may have lamellae, but the limitation of light microscopy inhibit the ability to view these features, and lamellae were also not often visible using SEM.

Three-dimensional shape Two and three-dimensional shapes are commonly used to describe and discriminate between starch morphotypes. In this study only three-dimensional (3D) shape was used as the granules were rolled during light microscopy to optimise visualisation of all planes of the granules. Fifteen different 3D shapes were observed within the comparative collection. These included a variety of rounded shapes such as spherical, sub-spherical, ovate, ellipsoidal, sub-ovate, pyriform (pear-shaped), reniform (kidney-shaped) and shapes with single or multiple facets such as dome, hemispherical, prismatic, polyhedral, quadrangular, cylindrical, conical, and prismatic. The nomenclature used to describe these shapes follow the International Code for Starch Nomenclature (ICSN) compiled in 2011 with some modification.

Rounded granules were observed within 27 of the 29 specimens within the comparative collection (see Table 5.2). The two species that did not have rounded starch morphotypes were D. esculenta and C. longa. The most commonly observed shape was spherical or “…a sphere in 66 which all radii are equal length.” (Reichert 1913), with a total of 22 specimens containing starch that matched this description. Within these specimens were members of a range of different families, and so there does not appear to be any diagnostic value in this shape for taxonomic classification. Likewise, sub-spherical granules were present within 15 specimens from the Anacardiaceae, Araceae, Convolvulaceae, Dioscoreaceae, Lecythidaceae, Moraceae, Musaceae, and Solanaceae. These are spherical granules with some small degree of curvature or are scalene, where all three main diameters are of unequal length. Also present in a large percentage of the reference collection were elongated granules which tend to be high in amylase (Gott et al. 2006). Ovate (n=16), subovate (n=2), and ellipsoidal (n=9) granules were commonly observed within members of the Dioscoreaceae and Musaceae families. These are distinguished from one another based on the diameter of the proximal, mesial and distal ends of the grain. Ellipsoidal granules have both distal and proximal ends equal in size, while ovate granules tend to be more egg-shaped with one end wider than the other. Sub-ovate are elongated granules with the same variation as described for sub-spherical granules. Other variations to these rounded granules included reniform or kidney-shaped granules, and pyriform or pear-shaped granules. These differ from the sub-ovate and sub-spherical granules based on a larger degree of curvature (reniform) and scalar width variation (pyriform). Pyriform granules were only recorded in those specimens belonging to the genus Musa, while reniform granules were present in five different specimens representing four different families including Dioscoreacae, Convolvulaceae, Musaceae, and Solanaceae.

More angular starch shapes were also very common within the comparative collection. Twenty eight of the 29 specimens had at least one angular morphotype. The only species that did not was Dioscorea pentaphylla. These shapes vary in the nature and number of straight or rounded sides (facets) that define the margins and diameters of the granules. Angular granules have been formed in compound starch granules, where a number of granules are clustered together within the parenchymatous cells, creating pressure facets. Single faceted 3D shapes include dome and hemispherical granules, which are differentiated from one another based on elongation. A domed granule is half an oval, while a hemispherical granule is half a sphere. Domed granules were present in 12 specimens, while hemispherical granules were slightly more common, being recorded within 15 specimens. Four of the five aroids, and both members of the Artocarpus genera within the comparative collection contained hemispherical granules. The remaining aroid, Xanthosoma sagittifolium, instead had dome-shaped granules. Some specimens contained both shapes, including Artocarpus heterophyllus, Amorphophallus paeoniifolius, the seeds of Barringtonia asiatica, Cyrtosperma merkusii, Morinda citrifolia and Piper methysticum.

Spherical Dome Reniform Amorphophallus paeoniifolius Ipomoea batatas Artocarpus heterophyllus Dioscorea nummularia Artocarpus altilis fruit Inocarpus fagifer seed Amorphophallus paeoniifolius Dioscorea pentaphylla Artocarpus altilis seed Ipomoea polpha Barringtonia asiatica fruit Ipomoea polpha Artocarpus heterophyllus Morinda citrifolia fruit Barringtonia asiatica seed Musa sp. 1 Barringtonia asiatica fruit Musa sp. 1 Cyrtosperma merkusii Solanum tuberosum Barringtonia asiatica seed Piper methysticum Horsfieldia palauensis Colocasia esculenta Pteridium sp. Ipomoea polpha Sub-ovate Cyrtosperma merkusii Spondias dulcis Morinda citrifolia fruit Musa sp. 1 Dioscorea nummularia Solanum tuberosum Musa sp. 1 Solanum tuberosum Dioscorea pentaphylla Tacca leontopetaloides Piper methysticum Horsfieldia palauensis Xanthosoma sagittifolium Pteridium sp. Xanthosoma sagittifolium Ovate Sub-spherical Alocasia macrorrhiza Dioscorea rotundata Alocasia macrorrhiza Inocarpus fagifer seed Amorphophallus paeoniifolius Horsfieldia palauensis Artocarpus altilis fruit Ipomoea polpha Artocarpus altilis fruit Musa sp. 1 Artocarpus heterophyllus Musa sp. 1 Artocarpus altilis seed Musa sp. 2 Barringtonia asiatica fruit Musa sp. 2 Dioscorea alata Piper methysticum Barringtonia asiatica seed Piper methysticum Dioscorea bulbifera Pteridium sp. Dioscorea bulbifera Spondias dulcis Dioscorea nummularia Spondias dulcis Dioscorea rotundata Solanum tuberosum Dioscorea pentaphylla Solanum tuberosum Ipomoea batatas Hemispherical Polyhedral Alocasia macrorrhiza Dioscorea bulbifera Alocasia macrorrhiza Horsfieldia palauensis Amorphophallus paeoniifolius Ipomoea batatas Amorphophallus paeoniifolius Ipomoea batatas Artocarpus altilis fruit Inocarpus fagifer seed Artocarpus altilis fruit Inocarpus fagifer seed Artocarpus altilis seed Morinda citrifolia fruit Artocarpus altilis seed Morinda citrifolia fruit Artocarpus heterophyllus Piper methysticum Artocarpus heterophyllus Pteridium sp. Barringtonia asiatica seed Spondias dulcis Colocasia esculenta Tacca leontopetaloides Colocasia esculenta Tacca leontopetaloides Cyrtosperma merkusii Xanthosoma sagittifolium Cyrtosperma merkusii Dioscorea esculenta Ellipsoidal Conical Quadrangular Pyriform Barringtonia asiatica fruit Curcuma longa Artocarpus altilis seed Musa sp. 1 Barringtonia asiatica seed Dioscorea alata Artocarpus heterophyllus Musa sp. 2 Dioscorea bulbifera Dioscorea bulbifera Dioscorea esculenta Dioscorea nummularia Dioscorea nummularia Ipomoea polpha Ipomoea polpha Dioscorea rotundata Cylindrical Prismatic Musa sp. 1 Musa sp. 1 Musa sp. 2 Xanthosoma sagittifolium Musa sp. 2 Musa sp. 2 Spondias dulcis Pteridium sp. Solanum tuberosum Tacca leontopetaloides

Table 5.2 Three-dimensional shapes of reference species

Multi-faceted angular shapes recorded included conical, cylindrical, polyhedral, quadrangular, and prismatic. A number of specimens contained starch that could be described as conical, having a flat circular base and a tapering top. These included Curcuma longa, most of the members of the Dioscorea genus apart from D. pentaphylla which has more ovate granules, both Musa spp., and Solanum tuberosum. Cylindrical granules fall part way between rounded and angular, being described as having “…a circular base and top, both of equivalent size.” (Reichert 1913). This shape was not commonly observed, and was present only in three specimens. Four specimens contained granules that had six sides, with four being elongated, and thus could be described as quadrangular. Within these four specimens, less than five of these granules were observed. The most common multi-faceted angular shape by a significant margin was polyhedral. Polyhedral granules are defined as “…having many faces that are not necessarily of the same two dimensional shape.” (Reichert 1913). These were recorded in all of the aroids, several members of the Dioscoreaceae family, and a range of other Monocots and Eudicots totalling 15 specimens. 68

Modifiers of shape These three-dimensional shapes can be altered by several other morphological attributes. Faceting was discussed above in terms of the development of overall shapes when granules are formed in compound clusters. Sometimes these ‘pressure facets’ are minimal, and only create slight impressions on the sides of granules. These granules are usually rounded in shape but have one or two pressure facets on the margins. Additionally, some dome or hemispherical granules can have an additional one to two pressure facets where these granules have been formed in clusters of three granules. They still retain the shape of half a sphere or oval. These types of pressure faceting are common in Amorphophallus paeoniifolius, Barringtonia asiatica, Cyrtosperma merkusii, Inocarpus fagifer, Ipomoea batatas, Ipomoea polpha, and Piper methysticum.

A small number of specimens within the comparative collection contained granules that were ‘compressed’ so that the granules had a smaller dimension in one plane than the other. These granules had to be consistently rolled to view this modification of shape in profile. Three of the yams- Dioscorea alata, Dioscorea bulbifera, and Dioscorea rotundata, and a ginger- Curcuma longa, all have conical or ovate starch granules that are frequently compressed. In addition, both Musa spp. within the comparative collection also have compressed forms, but these are less commonly observed than non-compressed granules. Equitorial grooves often coincide with compressed granules or those that have uneven dimensions in different planes, and can be seen as a fold that runs along the long axis of the granules when these are turned on edge (ICSN 2011). This feature was common in Dioscorea rotundata and noted in Artocarpus altilis, Amorphophallus paeoniifolius, Dioscorea alata and Piper methysticum.

Several shape modifications were restricted to only one or two particular specimens within the comparative collection, and so can be considered diagnostic attributes of starch morphotypes. The first of these were ‘projections’, which were defined as areas that extend beyond the main surface of the grain (Reichert 1913), and were present on all recorded granules from C. longa, and a small number of granules of Dioscorea pentaphylla. The conical starch granules deriving from these specimens also had sharply tapering tips that differentiated these from granules seen in other specimens. The starch of Morinda citrifolia were often noticeably ‘concave-convex’ where one side curves inward and the other curves outward (ICSN 2011), creating a hollowed appearance. Finally, the conical starch of Dioscorea bulbifera each had a distinctive ‘bend’ at the proximal end of the granule just below the hilum. This was sharper than a ‘curve’ which is defined as a smooth bend in the form of the granule (Reichert 1913).

Granule length and width The maximum length of starch granules is used by many starch analysts to identify unknown archaeological starch as a univariate statistical technique within an assemblage-type approach (Field et al. 2009; Therin et al. 1999). Following these studies, the range of granule lengths

69 within the comparative collection were analysed and compared as assemblages of each species (see Figure 5.3). Most of the recorded starch within the comparative collection falls into the 8- 25µm range (86% of species), and due to the relatively narrow range there is a significant amount of overlap between specimens. Only a small number of specimens have ranges that are larger or smaller than this. Several of the yams, Dioscorea pentaphylla and Dioscorea rotundata, as well as both Musa spp., Curcuma longa, Ipomoea polpha and Solanum tuberosum all have maximum lengths that could exceed 40µm. Of these, the Dioscorea spp. are the only specimens that have an average length above 40µm. Overall, Dioscorea pentaphylla has the largest starch within the comparative collection, with an average length of 90µm, and a maximum of 145µm and are therefore taxonomically distinctive. On the smaller end of the spectrum, several specimens have starch morphotypes that can range below 8µm. These include Artocarpus altilis seed and fruit, Artocarpus heterophyllus, Alocasia macrorrhiza, Colocasia esculenta, Cyrtosperma merkusii, Dioscorea bulbifera, Dioscorea esculenta, Horsfieldia palauensis, Inocarpus fagifer, Ipomoea batatas, Ipomoea polpha, Morinda citrifolia, Pteridium sp., Tacca leontopetaloides and Xanthosoma sagittifolium. The only specimen to have the entire range of maximum lengths below 8µm is C. esculenta, which has an average length of 4µm. Despite this, this species cannot be distinguished from others solely on this attribute, as a number of other specimens have overall ranges that overlap with Colocasia esculenta such as Alocasia macrorrhiza and Artocarpus altilis.

A study of granule widths revealed that there is substantial variation in the comparative collection based on this metric attribute. The starch of most taxa had widths between 7-19µm (93% of species), which is marginally smaller than the range of starch lengths for most species. However, unlike starch lengths, there is a greater number of specimens that contain starch smaller or larger than this range (see Figure 5.4). Sixteen specimens have granules that range in width above 19µm. Dioscorea pentaphylla is once again distinguishable by having the largest starch granules in this second dimension. A total of 20 specimens contain starch morphotypes that can range smaller than 7µm, but only six of these specimens have average widths below this point. The smallest width range was recorded within the starch of Colocasia esculenta which has a median of 4µm, and a total range of 2-7µm.

Clearly, there is a significant amount of overlap in these two variables in the comparative collection. Only one specimen can be confidently distinguished from others based on either of these variables. Dioscorea pentaphylla has starch that ranges larger in length and width that any other specimens. On the smaller end of the spectrums for both widths and lengths, C. esculenta consistently has the lowest medians for these attributes, but this species still overlaps with the total range of others within the comparative collection. Therefore it is not able to be distinguished solely on these attributes.

Other metric variables In addition to maximum length and width, a number of other metric variables were measured on all starch granules sampled from each species within the comparative collection. As mentioned previously, these included the ‘length of the extinction cross’, ‘maximum distance between the arms of the cross’, ‘distance between the hilum and the end of the granule’, and the ‘maximum angle within the arms of the extinction cross’. Most of these measurements are not diagnostic of species on its own when compared through univariate analyses of the starch assemblages in the comparative collection. These are more relevant when considered in multivariate statistical analyses as these dimensions are related to the three-dimensional shape and surface texture of starch granules. Therefore several of these will not be explored here, but instead will be compared as variables in the multivariate morphometric analysis later in this chapter.

Hilum position can be compared between specimens but only when this is considered as a ratio between the ‘length’ and the ‘distance between the hilum and the end of the granule’ (see Figure 5.5). This ratio enables statistical differentiation between starch that is centric (end-on) or eccentric (side-on). Centric hila are those that have a ratio that falls between 0.5-0.6, meaning that the hilum is close to the centre of the granule. These granules have been viewed and recorded end-on. Eccentric hila are those that have a ratio falling in the range of 0.61-1, and indicate that the granule has been viewed side-on. This is particularly relevant for granules that vary in dimension across different planes such as ovate or conical granules. When the distribution of these ranges is compared between specimens, it is clear that most sampled granules from each specimen have been recorded in both centric and eccentric views. However a number of specimens were primarily viewed side-on, including C. longa, D. alata, D. bulbifera, D. nummularia, D. pentaphylla, D. rotundata, Musa spp., S. tuberosum and S. dulcis. This is most likely a result of the ovate, sub-ovate or conical shape of these granules which makes them hard or impossible to view completely end-on. Another 18 specimens were primarily viewed end-on, and this is similarly a result of shape, but in this case granules are mostly spherical or polygonal and so are even in length across all dimensions. These hilum position ratios were used within the following multivariate statistical analysis to divide the starch within the comparative collection into comparable datasets.

Figure 5.3 Box plot of starch granule lengths within reference collection

Figure 5.4 Box plot of starch granule widths within reference collection

Figure 5.5 Box plot of starch granule hilum position to length ratios within reference collection

Length Width Length to Species Organ range range hilum position 3D shape Other features (µm) (µm) ratio range

Hemispherical, ovate, polygonal, Small vacuole and flat single and multiple faceting common. Artocarpus altilis Fruit 5-12 3-12 0.5-0.6 spherical, sub-spherical Equitorial grooves and lamellae rare.

Hemispherical, ovate, polygonal, Flat and curved single and multiple faceting common. Small vacuoles Artocarpus altilis Seed 3-12 3-9 0.5-0.75 quadrangular, spherical and longitudinal fissuring rare

Dome, hemispherical, polygonal, Artocarpus Flat and curved single and multiple faceting common. Small vacuoles Fruit 4-13 3-13 0.5-0.6 quadrangular, spherical, sub- heterophyllus and stellate or radial fissuring rare spherical Hemispherical, ovate, polygonal, Flat and curved single and multiple faceting common. Simple and Alocasia macrorrhiza Corm 2-10 2-9 0.5-0.6 sub-spherical compound grains. Flat and curved single and multiple faceting and longitudinal fissuring Amorphophallus Dome, hemispherical, ovate, Corm 12-36 7-30 0.5-0.71 or indentation common. Equitorial grooves rare. Simple and paeoniifolius polygonal, spherical compound grains. Dome, ellipsoidal, spherical, sub- Flat and curved multiple faceting common. Curved single faceting, Barringtonia asiatica Fruit 8-20 6-20 0.5-0.73 spherical large and small vacuoles and radial or transverse fissuring rare. Dome, ellipsoidal, Small vacuole and flat and curved single and multiple faceting Barringtonia asiatica Seed 8-20 6-17 0.5-0.78 hemispherical, spherical, sub- common. Large vacuole rare. spherical Hemispherical, polygonal, Flat multiple and single faceting common. Simple and compound Colocasia esculenta Corm 3-8 2-7 0.5-0.75 spherical, sub-spherical grains. Curcuma longa Rhizome 15-60 14-30 0.8-1 Conical All lenticular, lamellae, proximal protrusion and tapered tip. Small vacuole and flat and curved single and multiple faceting Dome, hemispherical, polygonal, Cyrtosperma merkusii Corm 5-26 5-26 0.5-0.6 common. Radial or irregular fissuring and large vacuole rare. Simple spherical and compound grains. Commonly lenticular and lamellae present. Equitorial groove, small Dioscorea alata Root tuber 10-25 5-19 0.62-1 Conical, ovate vacuole and radial fissuring rare. Conical, hemispherical, Commonly lenticular or wedge-shaped, bent with flat multiple and Dioscorea bulbifera Aerial bulbil 5-30 4-26 0.62-0.96 ellipsoidal, ovate, sub-spherical, single faceting and lamellae present and rare radial fissuring. triangular Flat and curved multiple and single faceting common. Simple and Dioscorea esculenta Root tuber 3-13 3-11 0.5-0.75 Polygonal, quadrangular compound grains. Dioscorea Conical, ellipsoidal, kidney, Lamellae and single flat faceting common. Multiple flat faceting, small Root tuber 11-50 10-39 0.62-0.9 nummularia ovate, spherical vacuole and longitudinal fissuring rare.

Table 5.3 Summary of starch morphology within reference collection 74

Length Width Length to Species Organ range range hilum position 3D shape Other features (µm) (µm) ratio range Lamellae present. Commonly lenticular with equitorial groove and Dioscorea rotundata Root tuber 34-95 15-62 0.62-0.94 Conical, ovate, sub-spherical parallel fissuring. Small vacuole rare. Horsfieldia Dome, ovate, polygonal, Small vacuole and flat multiple and single faceting common. Lamellae Nut 5-27 4-24 0.5-0.67 palauensis spherical rare. Hemispherical, polygonal, Small vacuole and flat multiple and single faceting common. Inocarpus fagifer Seed 6-21 6-19 0.5-0.6 spherical, sub-spherical Longitudinal fissuring or indentation rare. Hemispherical, polygonal, Small vacuole and flat and curved multiple and single faceting Ipomoea batatas Root tuber 4-17 4-16 0.5-0.6 spherical, sub-spherical common. Lamellae and transverse fissuring rare. Dome, ellipsoidal, Small vacuole and stellate, transverse, radial or irregular fissuring hemispherical, kidney, Ipomoea polpha Root tuber 7-51 7-46 0.5-0.73 common. Flat and curved multiple and single faceting, lamellae and polygonal, quadrangular, large vacuoles rare. spherical, sub-spherical Dome, hemispherical, polygonal, Large vacuole, concave-convex and curved multiple and single faceting Morinda citrifolia Fruit 5-15 4-14 0.5-0.6 spherical common. Flat multiple and single faceting rare. Conical, dome, ellipsoidal, kidney, ovate, pyriform, Musa sp. 1 Fruit 9-53 7-32 0.62-0.95 Lamellae, flat single faceting and lenticular shape rare. spherical, sub-ovate, sub- spherical Conical, cylindrical, ellipsoidal, Lamellae and small vacuole common. Flat single faceting, longitudinal Musa sp. 2 Fruit 8-61 6-48 0.62-0.92 ovate, pyriform, sub-spherical fissuring and lenticular shape rare. Small vacuole present. Lamellae and longitudinal, stellate, irregular, Dome, hemispherical, ovate, Piper methysticum Primary root 8-26 4-20 0.5-0.6 radial or mesial cleft fissuring common. Flat and curved multiple and spherical, sub-spherical single faceting, lenticular shape, and equitorial groove rare. Dome, ellipsoidal, ovate, Flat and curved single faceting common. Flat and curved multiple Pteridium sp. Rhizome 4-13 3-11 0.5-0.6 polygonal, spherical faceting and small vacuole rare. Conical, kidney, ovate, spherical, Lamellae present. Small vacuole common. Flat single faceting and Solanum tuberosum Stem tuber 11-62 10-42 0.65-0.89 sub-ovate, sub-spherical oblique fissuring rare. Cylindrical, hemispherical, Small or large vacuole present. Lamellae and stellate, branching or Spondias dulcis Fruit 13-32 8-27 0.5-0.94 ovate, spherical, subspherical transverse fissuring common. Flat single faceting rare. Conical, cylindrical, Tacca Flat and curved multiple and single faceting common. Longitudinal Stem tuber 4-20 3-17 0.5-0.67 hemispherical, polygonal, leontopetaloides fissuring rare. Simple and compound grains. spherical Xanthosoma Dome, polygonal, prismatic, Flat and curved multiple and single faceting common. Longitudinal Corm 4-13 3-12 0.5-0.6 sagittifolium spherical fissuring rare. Simple and compound grains.

Multivariate statistical analysis of starch Discriminant Function Analysis Multivariate statistics in the form of Discriminant Function Analysis (DFA) was used to discriminate between starch morphotypes observed at various taxonomic levels within the comparative collection used for this study. DFA is a form of linear discrimination where classification is based on the probabilities of an individual granule coming from groups with particular means and variances (Torrence et al. 2004). This method is naturally comparative, where data regarding modern plant taxa for which the taxonomy is known, provides the basis for making predictions about unknown or unclassified individuals (Kovarovic et al. 2011), making this statistical technique ideal for morphometric comparisons here. DFA utilises predictor variables to determine the linear dimensions along which known groups are best separated (Tabachnick and Lidell 2007). A new set of variables, known as discriminant functions, are derived from building linear combinations of the original variables that maximise between within-group variance. The number of discriminant functions is equal to the number of groups minus one, and unless there are fewer predictor variables than groups, they only account for the variance which best discriminates groups according to that model. Other possible options such as logistical discrimination and nearest neighbour make fewer or no assumptions of the data, but do not allow ready interpretation of the results in terms of individual variables (Drennan 1996; Torrence et al. 2004), which is important for this study.

Data was collected during light microscopy and subsequent image analysis, whereby 18 attributes of 50 granules per sample were recorded, formed the dataset for the DFA. Each grain had been rolled to view the 3D shape, but the metric variables including length and width were measured off a photograph taken at a particular orientation. The Excel reference database was therefore divided into two based on the orientation or view (eccentric/side-on or centric/end-on) through which each grain had been recorded.

View Metric Binary (presence or absence) Centric Length 3D Shape- dome/hemispherical Small vacuole at hilum Width 3D Shape- ellipsoidal/ovate Multiple facetting Length of cross 3D Shape- reniform Flat facetting 3D Shape- angular (polyhedral, prismatic, Maximum distance between arms of cross Curved/rounded facetting quadrangular, hexagonal) Maximum distance between hilum and end 3D shape- pyriform/subspherical Fissuring at hilum Maximum angle of the cross 3D Shape- spherical Concave-convex Equitorial groove Large vacuole at hilum Lamellae

Eccentric Length 3D Shape- conical/triangular Lamellae Width 3D Shape-cylindrical Vacuole at hilum Length of cross 3D Shape- dome/hemispherical Multiple facetting Maximum distance between arms of cross 3D Shape- ellipsoidal/ovoidal Flat facetting Maximum distance between hilum and end 3D Shape- reniform Curved/rounded facetting 3D Shape- angular (polyhedral, prismatic, Maximum angle of the cross Fissuring at hilum quadrangular, hexagonal) 3D Shape- spherical Large vacuole 3D Shape- subspherical/subovoidal Tapered tip Compressed Bent Equitorial groove

Table 5.4 Description of metric and binary variables used during Discriminant Function Analysis 76

In order to be able to combine the data recorded from the range of ordinal, binary and nominal variables, the nominal (or discontinuous) variables were converted into a series of binary attributes. Thus 3D shape was converted into 14 binary variables, and the style of the extinction cross was converted into three binary variables, each indicating the presence or absence of that particular shape or style for that grain. After this data manipulation, a total of 34 variables were included in the initial DFA datasets and outputs.

PAST statistical software was used to conduct the multivariate statistical analyses. To set a baseline for subsequent dataset comparison, both reference databases were initially combined and entered into the PAST software together. The DFA was run using linear discrimination and a confusion matrix was constructed based on the ‘classifier’ output from this function. Essentially the DFA used the attributes included in the dataset as a ‘learning sample’ to predict the classification of each starch grain, and to establish how well each species could be re-classified back into its own grouping. The confusion matrix gave an overall percentage of correct re-classifications for the dataset, which when both orientations were combined was 48.3%.

This figure is statistically quite low, and emphasised the need to attempt the division of the reference collection according to orientation, so that starch would only be compared to other granules recorded in the same view. Due to the three dimensional nature of starch granules, granules can be imaged and recorded from many different angles. In order to establish some consistency within the datasets, the orientation was decided according to an acceptable range of length to hilum position ratios for eccentric and centric views. As discussed earlier, this ratio was calculated for each reference starch grain by dividing the maximum distance between the hilum and the end of the grain, by the total length of the grain. A centric or end-on grain could have a ratio of 0.5-0.6, and an eccentric or side-on grain could have a ratio of 0.61-1.0. The database was thus divided according to these ratio ranges. Each reference database was entered into the software separately, and outputs created in the same manner as the combined dataset. Within the eccentric (side-on) view, the confusion matrix calculated that the total percentage of correct re-classifications for that view was 53.6%. The centric dataset had a total percentage of 34%.

Clearly, the starch morphotypes in the eccentric dataset are more easily discriminated from each other than those in the centric dataset. To increase the percentage of correct re- classifications within the centric dataset and thus ensure that the archaeological starch can be more confidently classified using DFA, some changes were made to the variables included in the datasets. The ‘loadings’ which outline the role of each variable towards discrimination within the DFA for each dataset were reviewed, and decisions were made to remove or re- arrange some binary categories. The DFA was run with these new datasets, and higher

77 percentages of successful reclassification of species were achieved within both the eccentric and centric views. The final DFA were run based on these revised datasets. Overall, the percentage of starch granules within the eccentric dataset that were correctly reclassified to species was 72.98%. Understanding these values was vital towards characterizing starch morphology at species level. Plots were then created that visually demonstrated the ability to discriminate between each of the species in each of the datasets (see Figure 5.6and Figure 5.7). . These also indicated the amount of discrimination due to each of the first two canonical variates, which form the axis for the plot.

Eccentric Centric Amorphophallus paeoniifolius Alocasia macrorrhiza Artocarpus altilis Amorphophallus paeoniifolius Barringtonia asiatica Artocarpus altilis Colocasia esculenta Barringtonia asiatica Curcuma longa Colocasia esculenta Dioscorea alata Cytrosperma merkusii Dioscorea bulbifera Dioscorea esculenta Dioscorea esculenta Inocarpus fagifer Dioscorea nummularia Ipomoea batatas Dioscorea pentaphylla Morinda citrifolia Musa spp. Piper methysticum Solanum tuberosum Pteridium sp. Spondias dulcis Spondias dulcis Tacca leontopetaloides Tacca leontopetaloides Xanthosoma sagittifolium

Table 5.5 Species included in centric and eccentric datasets for Tongan analysis

The reference collection up until this point had included a number of species that have never been recorded in Tonga, such as Ipomoea polpha and Dioscorea rotundata but which had been included to further our understanding of starch morphology at genus level within the larger comparative collection. Ultimately the point of this analysis is to classify archaeological starch with a high degree of confidence, and the inclusion of these species was making the discrimination between species in both views more difficult due to similarities between their recorded starch morphology. Species were therefore restricted to those found in Tonga in late prehistory and also common contaminants. The final species list to be used in Part Two of this thesis for the identification of archaeological starch included fourteen species within the eccentric dataset, and fifteen species in the centric dataset, but again these were not necessarily the same species within both datasets. The overall success rates of correct reclassifications within these new datasets were 68.4% for eccentric, and 56.7% for centric.

Figure 5.6 Plot showing discrimination of species within centric dataset according to first two canonical variates

Figure 5.7 Plot showing discrimination of 15 species within eccentric dataset according to first two canonical variates

Classification matrices When the confusion matrices for each of the datasets are analysed, it becomes apparent that some species contain starch morphotypes that are more distinctive than others. In the centric dataset only eight of the 20 specimens had over 60% of granules correctly re-assigned. The

79 specimen with the highest percentage of correct re-assignments were Colocasia esculenta with 77%, followed closely by Inocarpus fagifer with 76%, Morinda citrifolia and Spondias dulcis with 74%, Amorphophallus paeoniifolius with 72%, and Pteridium sp. with 71%. The specimens that were the least able to be differentiated within the eccentric view (i.e. under 30%) include Barringtonia asiatica seed (11%), Xanthosoma sagittifolium (24%), Artocarpus heterophyllus (25%), and Artocarpus altilis seed (26%). As discussed previously, starch from several species was more easily distinguished within the eccentric view, and 12 of the 19 specimens included in this dataset had over 60% of granules correctly re-assigned. Of these, two specimens, C. esculenta and S. dulcis, had all recorded starch re-assigned to the original specimens. Only slightly less successful than these was Curcuma longa with 98% and Dioscorea rotundata with 94%. The specimens with the lowest percentages of correct re- assignment within the eccentric dataset, and thus the least able to be differentiated using attributes within the dataset are Dioscorea esculenta (24%) and Ipomoea polpha (30%).

Automated classification of granules of unknown origin The multivariate statistical analysis fulfilled several goals within the morphological analysis of the comparative collection. Firstly, the analysis emphasised those attributes that were most useful towards discriminating between the starch morphotypes found in specimens included in the comparative collection. Of the 21 variables compared within the centric dataset, and the 25 variables in the eccentric dataset, the metric variables consistently had the highest loadings. This suggests that these attributes are an important source of morphological variation. Of these, starch ‘length’ had the highest loading in the centric dataset, while ‘length of cross’ and ‘maximum distance between the arms of the cross’ had higher loadings within the first two canonical variates in the eccentric dataset. Surprisingly all other attributes, such as categories of three-dimensional shapes, contributed minimally to the differentiation of species within the comparative collection which implies that these should not be heavily relied on for identification of unknown granules. The automated classification of unknown granules using DFA is a useful first step, but should be expressed with varying levels of confidence and visually cross-checked before final confirmation of taxonomic identifications, as will be explored in Part Two of this thesis.

Figure 5.8 Classification matrix for the overall centric dataset, showing highest discrimination of Colocasia esculenta, Inocarpus fagifer, Morinda citrifolia and Spondias dulcis (species listed vertically in the first column are the original species, and those listed horizontally in the top row are the species to which DFA classified granules)

Figure 5.9 Classification matrix for the overall eccentric dataset, showing highest discrimination of Colocasia esculenta, Curcuma longa, and Dioscorea pentaphylla (species listed vertically in the first column are the original species, and those listed horizontally in the top row are the species to which DFA classified granules)

Morphology of vegetative storage parenchyma

Morphological analysis of fresh samples Hather (1991, 2000) discriminates between the vegetative storage parenchyma that is found in roots and tubers, and all other parenchymatous organs such as fruits, seeds, and nuts. He makes this distinction because the function of these organs is to store reserve starch and carbohydrates during photosynthesis for plants, and thus these organs share many similar morphological traits. It is this storage function, and subsequent calorific and nutritional value, that led to the development of many prehistoric subsistence strategies based upon the collection or cultivation of these starchy organs (Barton and Paz 2007; Denham 2007b; Golson 2007; Holden, Hather and Watson 1995).

In the comparative collection of modern parenchyma developed for this study, a number of fruits and seeds were also included. The justification for these inclusions was that these organs also often produce starch and other vitamins and minerals, and have been recorded in the ethnographic record for Tonga (Beaglehole and Beaglehole 1941; Gifford 1929; Whistler 2009) to supplement diets either on a regular basis or during times of famine. Fruits such as breadfruit (Artocarpus altilis), the tropical almond (Inocarpus fagifer), and bananas (Musa spp.) were grown bordering plantations of yams (Dioscorea spp.), aroids (Colocasia esculenta, Alocasia macrorrhiza, Cyrtosperma merkusii), sweet potato (Ipomoea batatas), kava (Piper methysticum) and arrowroot (Tacca leontopetaloides). Under the premise that these crops were being processed and cooked in the same areas as the roots and tubers, it is possible that some macro-remains of these organs may end up charred in hearths and were preserved in archaeological contexts. This has been documented elsewhere in Polynesia by the recent find of breadfruit in Tahiti (Kahn and Ragone 2013).

The organisation of tissues within fruits does differ from that of stem and root-derived organs, and therefore they were described separately. The basic cell morphology and types of boundary, ground and some vascular tissues were recorded for each sample of non-vegetative storage parenchyma. Some similarities emerged such as the nature of the ground tissues and the arrangement of vascular bundles where these were present within the organ. Most of the fruits included in this study are Dicotyledons or Eudicotyledons, and thus have different anatomical features from Monocotyledons which form the bulk of the species within the comparative collection. This data thus provided a means of comparison with the stem and root-derived organs. Here, fresh samples will be analysed using univariate and multivariate statistical analyses to study parenchyma morphology, with a small study upon the changes that occur during charring. These studies summarise the descriptions provided in Appendix A.

Ground tissue morphology Some of the distinguishing features of parenchyma ground tissue are best described through comparison of these tissues with hardwood. Some differences to wood charcoal include consistent cell shapes that are usually rounded or angular; the presence of distinctive vascular bundles or tissues, and there are usually very few rays dividing these tissues. When comparing vegetative and non-vegetative storage parenchyma from within roots, tubers, fruits and seeds with each other these morphological features need to be teased apart to allow identification of charred material where possible.

As described above, 40 cells from each specimen were recorded to assess the morphology of parenchymatous ground tissues within roots, tubers, fruits and seeds. The length and width of each cell was recorded, along with cell shape (rounded, angular or rounded- irregular), and cell dimensions (isodiametric or elongated). On top of these attributes of the individual cells, some overall descriptions were made about the ground tissue. These included the overall presence or absence of inter-cellular spaces, and the variety of cell contents that had been preserved in the histological thin sections.

Most species contained ground tissue that were consistently either rounded (60%, n=26), rounded-irregular (12%, n=5), or angular (21%, n=9); however a small percentage (7%, n=3) had both rounded and angular cell shapes within the same specimen. These included Artocarpus altilis, Colocasia esculenta, and Dioscorea alata, and interestingly these species do not share any phenotypical or genotypical traits with one another aside from the fact that all three specimens are types of starchy organs. These specimens represent a range of different storage organs including a fruit (A. altilis), a corm (C. esculenta) and a tuber (D. alata). The yams (Dioscoreaceae) are traditionally classified as root tubers; however the anatomy of these organs is much closer to stem-derived tissue than true roots. Onwueme (1978) argues that the yam tuber is “probably neither a root nor a stem structure. Rather it is a structure that originates from the hypocotyls- the transition zone between the stem and the root”. With this in mind, the attributes for describing stem anatomy have been used here for yams rather than root anatomy, especially with regard to the arrangement of vascular tissues and overall tissue organisation.

The ground tissues of the remaining yams (Dioscorea bulbifera, Dioscorea esculenta, and Dioscorea nummularia) and the Polynesian arrowroot (Tacca leontopetaloides), which also belongs to the Dioscoreaceae family, are all rounded in shape. Similarly, the remaining aroids (Alocasia macrorrhiza, Cyrtosperma merkusii, and Xanthosoma sagittifolium) have rounded cells within the ground tissues. Other starchy crops such as the bananas (Musa spp.), kava (P. methysticum) and a type of ginger (Zingiberaceae sp.) contain more irregularly rounded cells within the ground tissue. The majority of the Pteriodophytes or ferns have more angular shaped cells, including Asplenium sp., Pteridium sp., and Todea sp. Similarly many of the fruits such as 83

Barringtonia asiatica, Barringtonia racemosa, Ficus tinctorius, Spondias dulcis, and Syzygium malaccense have angular ground tissue cells.

Rounded Rounded- irregular Angular Artocarpus altilis seed Epipremnum pinnatum Musa sp. 1 Asplenium sp. Alocasia macrorrhiza Ficus copiosa Musa sp. 2 Barringtonia asiatica fruit Angiopteris sp. Inocarpus fagifer Piper methysticum Barringtonia racemosa seed Asplenium sp. Morinda citrifolia Solanum tuberosum Ficus tinctorius Araceae sp. Pangium edule Zingiberaceae sp. Ipomoea batatas Barringtonia asiatica Pueraria lobata Pteridium sp. seed Barringtonia racemosa Saccharum officinarum Spondias dulcis fruit Cordyline fruiticosa Pandanus tectorius Mixed Syzygium malaccense Tabernaemontana Cyrtosperma merkusii Artocarpus altilis fruit Todea sp. aurantiaca Dioscorea bulbifera Tacca leontopetaloides Colocasia esculenta Dioscorea esculenta Xanthosoma sagittifolium Dioscorea alata Dioscorea nummularia Zingiberaceae sp.

Table 5.6 Ground tissue cell shapes of taxa in reference collection

The majority of specimens included in this analysis had ground tissue parenchyma cells that were a mixture of elongated (greater in length than width) and broadly isodiametric (roughly even in length and width) in dimension. These specimens with mixed dimensions totalled 37 out of the full 43 specimens, equating to 86%. The remaining 14% (n=6) of specimens had consistently broadly isodiametric cell dimensions. These included Barringtonia asiatica fruit, Barringtonia racemosa seed, Dioscorea nummularia, Ficus copiosa, Saccharum officinarum, and Xanthosoma sagittifolium, and represent a range of different plant families and organ types. There does not appear to be any patterning that may suggest why this particular range of specimens has solely isodiametric cells within the recorded random sample of ground tissue recorded.

Isodiametric Mixed Barringtonia asiatica fruit Artocarpus altilis seed Ipomoea batatas Barringtonia racemosa seed Artocarpus altilis fruit Inocarpus fagifer Dioscorea nummularia Alocasia macrorrhiza Morinda citrifolia Ficus copiosa Angiopteris sp. Musa sp. 1 Saccharum officinarum Asplenium sp. Musa sp. 2 Xanthosoma sagittifolium Araceae sp. Pangium edule Barringtonia asiatica seed Pueraria lobata Barringtonia racemosa fruit Piper methysticum Colocasia esculenta Pandanus tectorius Cordyline fruiticosa Pteridium sp. Cyrtosperma merkusii Spondias dulcis Dioscorea alata Syzygium malaccense Dioscorea bulbifera Solanum tuberosum Dioscorea esculenta Tabernaemontana aurantiaca Dioscorea nummularia Tacca leontopetaloides Epipremnum pinnatum Todea sp. Ficus tinctorius Zingiberaceae sp.

Table 5.7 Ground tissue cell dimensions of taxa in the reference collection 84

The statistical analysis of cell lengths and widths points to a large degree of overlap in these particular attributes of ground tissue morphology. Box plots were created using the PAST statistical software to visually display the range of cell lengths and widths for each of the specimens included in the modern comparative collection. Outliers were indicated within the ranges of almost every species and organ, indicating that the random sample of 40 cells recorded in this study may not fully represent the degree of variation within the ground tissue of each specimen.

Despite this, the box plots demonstrate that most cell lengths within the comparative collection fall into a range larger than 40µm and smaller than 160µm. A number of species have cell lengths that could exceed 160µm, including Alocasia macrorrhiza, Angiopteris sp., Asplenium sp., Dioscorea alata, Dioscorea esculenta, Dioscorea nummularia, Ficus copiosa, Ficus tinctorius, Ipomoea batatas, Musa sp. 2, Spondias dulcis, Solanum tuberosum, Tacca leontopetaloides and Todea sp. The overall ranges of many of these species are very similar, and therefore the diagnostic value of this attribute of ground tissue morphology is reduced. To narrow this down further it may be more useful to consider the smaller number of species that have cell lengths that can exceed 240µm. These include Angiopteris sp., Asplenium sp., D. esculenta, D. nummularia and S. dulcis. The largest cells recorded within the comparative collection are in the ground tissues of Asplenium sp. and D. esculenta, which both can exceed 320µm in length. On the smaller end of the spectrum, some specimens have cell lengths that can range below 40µm such as the fruit of Artocarpus altilis and the stem of Saccharum officinarum. The two samples of the fruit phalange of Pandanus tectorius, and the root tuber of D. nummularia indicate that there can be some intra-species diversity in cell lengths, and therefore this attribute of ground tissue morphology should be considered mostly undiagnostic on its own.

All taxa within the reference collection have cell widths within the ground tissues that can range between 30-120µm. However, just over a third of the specimens have cell width ranges where the minimum widths are smaller than 30µm (37%, n=16). These include A. altilis fruit, Araceae sp., B. asiatica fruit, B. racemosa seed, C. merkusii, F. copiosa, F. tinctorius, I. fagifer, M. citrifolia, Musa sp. 1., Pangium edule, S. dulcis, Syzygium malaccense, S. officinarum, and both Zingiber spp. The specimens that have ground tissue cell widths that range above 120µm (35%, n=15) include Alocasia macrorrhiza, Angiopteris sp., Asplenium sp., D. alata, D. bulbifera, D. esculenta, D. nummularia, F. tinctorius, Musa sp.2, S. dulcis, S. malaccense, S. tuberosum, T. leontopetaloides and Todea sp.. The largest cell widths by quite a significant margin are those of D. esculenta, which range up to 230µm and so is a diagnostic attribute of parenchyma from this species.

Considering the degree of overlap between specimens in the univariate analysis of each of the recorded metric attributes, it was prudent to carry out a multivariate statistical study that could assess the relationships between sets of attributes in the modern comparative collection. A dataset was created in the PAST program that included the data of four attributes: cell lengths, widths, shapes and dimension of 40 recorded ground tissue cells from each specimen. Cell shape and dimensions were turned into nominal variables, with a number from 1-3 representing each shape (rounded, rounded-irregular or angular) and 1-2 representing each type of dimension (isodiametric or elongate).

DFA of the parenchyma attributes was carried out in the same way that the starch samples were analysed. A confusion matrix was created, with a total of only 21.8% of the recorded cells being correctly re-classified back into the original species. This figure is statistically very low, indicating that the majority of modern specimens could not be easily disciminated from one another based on these four attributes. Despite this, a small number of specimens were able to be separated from the remaining groupings reasonably well. The most easily discriminated was Musa sp. 1,which had a total of 78% correct reclassifications into original species. Closely following this was Dioscorea esculenta with 73%. Several species including the seed of Barringtonia racemosa (60%), and the fruit of Ficus copiosa (58%) had over half of all the cells correctly reclassified. The outputs of the DFA complemented the outcome of the univariate analysis, suggesting that there is substantial overlap within the morphology of ground tissues of many of the specimens in the modern comparative collection. Clearly some species will be more easily identified during the sorting of archaeological samples based on the morphology of ground tissue, such as those that have cells larger than 160µm or smaller than 40µm, those that have mixed cell shapes or consistently isodiametric cell dimensions. The majority of cells do not fall within these ranges, and so other non-metric attributes need to also be considered.

Figure 5.10 Box plot of parenchyma cell lengths of taxa in the reference collection 87

Figure 5.11 Box plot of parenchyma cell widths of taxa in the reference collection 88

Figure 5.12 Plot showing classification of parenchyma within reference collection using DFA

Other features of the ground tissue can include the presence of inter-cellular spaces, sclerenchyma, collenchyma, fibre bundles, duct cavities and cell contents. These can also be used to differentiate some species from one another. For example, both Musa sp. 1 and Musa sp 2. have duct cavities three to four cells long throughout the ground tissue that separate rows of parenchymatous cells. These species also contain fibre bundles which are also common throughout Zingiber spp., Asplenium sp., Cordyline fruticosa, Epipremnum pinnatum, and Pandanus tectorius. Regions of sclerenchyma can be observed within the pith of the fruit of Artocarpus altilis, Pangium edule, and Piper methysticum; while collenchmya forms the pith of Angiopteris sp. Inter-cellular spaces are present in most ground tissues; however it is absent in Dioscorea bulbifera, Pteridium sp., Spondias dulcis, Saccharum officinarum, and Solanum tuberosum. Cell contents can include also raphides, starch granules, druses and other types of crystals. Raphides are most commonly seen in aroids, but are also noticeable in the ground tissue of Tacca leontopetaloides.

Vascular tissue morphology The overall tissue organisation within the parenchyma of the modern specimens varies between the root and stem-derived organs, and the fruits and seeds, alongside the typical anatomical differences between Monocots and Dicots. The vascular tissues within these types of organs therefore also vary. Bundles of vascular tissues are present in stem-derived organs, where the phloem and xylem are either encircling or abutting one another. These bundles can be organised within the organ in one of three ways: 1) Dictyostele, where the bundle is formed as a chamber surrounding the pith and can be broken into segments; 2) Eustele, where bundles are arranged concentrically as either primary or secondary tissues radiating from the pith; or 3) Atactostele, where the bundles are organised seemingly haphazardly within the ground tissue of the organ. 89

The arrangement of the xylem and the phloem within the bundles can also be categorised into morphological types. These include:

 Amphivasal concentric— xylem completely encircles the phloem;  Amphivasal open ends— xylem almost completely encircles phloem apart from each end of the bundle;  Amphivasal u-shaped-—xylem partially surrounds the phloem in a u-shape formation;  Amphicribal concentric— phloem completely encircles the xylem;  Amphicribal open ends— phloem almost completely encircles xylem apart from each end of the bundle;  Amphicribal u-shaped— phloem partially surrounds the xylem in a u-shape formation  Open collateral- bundles of phloem and xylem abut one another with a region of cambium between the two types of tissue;  Closed collateral- bundles of phloem and xylem abut one another with no region of cambium between the two types of tissue;  Bicollateral—bundles of phloem and xylem abut one another with a region of cambium between the two types of tissue, and another additional bundle of phloem or xylem is present below this arrangement.

Figure 5.13 Description of vascular bundle arrangements within vegetative parenchyma (from Hather 2000)

When considering the stem-derived organs such as corms, rhizomes and stem tubers, it is clear that the majority of vascular tissues are organised within an atactotstele morphology (58%, n=11). Much smaller numbers are organised within a eustele (16%, n=3), or dictyostele arrangement (26%, n=5). Those specimens that contain an atactostele arrangement of vascular tissues tend to be corms and stem tubers of Monocots, while the dictyostele arrangement is most 90 common in the Pteriodophytes or fern rhizomes, and the eustele arrangement is commonly seen in Dicots. When the arrangements of vascular tissues within these bundles are broken down into their respective categories, further patterning becomes apparent. Some specimens did not have visible or clear vascular tissues and so were not included in the below table (see Table 5.8).

All of the aroids (Araceae) assessed within this study have vascular bundles within an atactostele pattern of stele organisation (as these are Monocots), and the vascular tissues are of an amphivasal arrangement where the phloem is surrounded by the xylem. Colocasia esculenta is the only aroid to contain two different types of amphivasal bundling, having both amphivasal concentric and u-shaped. Both Alocasia macrorrhiza and Cyrtosperma merkusii have amphivasal open-ended bundles, while Xanthosoma sagittifolium has bundles of amphivasal concentric arrangement. Another family belonging to the Monocots are the yams (Dioscoreaceae) and therefore have vascular bundles within an atactostele organisation; however these tissues differ from those belonging to the aroids as they are all of collateral bundling arrangement. Dioscorea alata and both samples of Dioscorea nummularia have bundles of open collateral arrangement with a region of cambium between the vascular tissues, while Dioscorea esculenta has closed collateral bundles without the layer of cambium. The only species from Dioscoreaceae to have a bicollateral arrangement is Tacca leontopetaloides, but outside this family Solanum tuberosum and Saccharum officinarum also contain bundles of this arrangement. The two specimens belonging to the Zingiberaceae family both have vascular tissues of amphicribal concentric arrangement where the xylem is surrounded the phloem.

Stem- Atactostele Collateral Amphivasal Amphicribal Dioscorea alata Alocasia macrorrhiza Zingiberaceae sp. Dioscorea bulbifera Colocasia esculenta Dioscorea esculenta Cordyline fruticosa Stem- Dictyostele Dioscorea nummularia Cyrtosperma merkusii Angiopteris sp. Tacca leontopetaloides Xanthosoma sagittifolium Asplenium sp. Solanum tuberosum Epipremnum pinnatum Pteridium sp. Saccharum officinarum Todea sp. Fruit Collateral Amphicribal Unknown Ficus copiosa Artocarpus altilis Pangium edule Ficus copiosa Morinda citrifolia Amphivasal Inocarpus fagifer Ficus tinctorius Barringtonia asiatica Musa sp. 1 Barringtonia racemosa Musa sp. 2 Root Pandanus tectorius Spondias dulcis Ipomoea batatas Syzygium malaccense Pueraria lobata Tabernaemontana aurantiaca Piper methysticum

Table 5.8 Vascular tissue arrangements of taxa in reference collection

The arrangement of vascular tissues within the fruits can vary significantly between samples. Where vascular bundles are present, these are not organised within an atactostele, eustele or dictyostele arrangements, but depend on other anatomical features of the organs. The arrangement of the tissues in the bundles is described, but not the overall organisation within the fruit as these do not fit any of the morphological categories used previously. The main purpose of including these samples is the description of basic cell and vascular morphology, in order to explore some differences between these organs and vegetative storage parenchyma. Almost half of the fruit specimens included within the comparative collection have vascular bundles of amphicribal arrangement (47%, n=8). Within these are a number of specimens with amphicribal concentric bundles including both Musa spp. and Tabernaemontana aurantiaca, and one example of a specimen with amphicribal u-shaped in Artocarpus altilis. A different version of amphicribal arrangement is seen in Syzygium malaccense where a stellate pattern of the xylem is present. Both fruits of Barringtonia spp. and Pandanus tectorius contained amphivasal concentric bundles. Finally, there is one specimen that contains vascular tissues with more than one type of arrangement. Ficus copiosa has amphicribal concentric and u-shaped alongside closed collateral bundles.

The true roots have vascular tissues that are regions of parenchyma within the stele and cortex of the organs, and there is very little variation between Monocotyledons and Dicotyledons. In primary root tissues, the cortex is often wide and the stele is central with very little pith (Hather 2000:61). A region of endodermis and pericycle separate the cortex from the stele, within which are alternate regions of phloem, xylem and cambium. The number of regions of vascular tissues within the stele can be as few as two (diarch), three (triarch), four (tetrarch), five (pentarch) or many (polyarch). Secondary root tissues have a different organisation of the vascular tissues, where the cortex is shed and the stele expands to replace the ground tissue by expanding centripetally to produce xylem and centrifugally to produce phloem and parenchyma. The centre of the organ is therefore composed of xylem and the outer region is composed of phloem and pericyclic parenchyma with a cambium separating these, and a periderm on the external surface.

The root samples within the modern comparative collection have some variation within the vascular tissues. The specimen of Pueraria lobata or Kudzu is a primary root tissue and has a polyarch xylem organisation within the stele with many vessels. Kava (Piper methysticum) root is a secondary structure that has many arms of fibre and xylem radiating from the pith, and bundles of phloem in between these. Sweet potato or Ipomoea batatas has a typical vesicular xylem that shows signs of anomalous tertiary growth next to vessels, surrounded by parenchymatous phloem outside the cambium.

Having considered the nature of bundle arrangements, it became clear that a level of overlap within the morphology of these vascular tissues exists. A univariate study of bundle lengths was carried out to assess the potential role of this attribute in distinguishing between species within the comparative collection. As demonstrated in the box and whisker plot (below, see Figure 5.14), there is again significant overlap in the measured specimens, with the majority of specimens falling into the 150-600µm range. However a number of species are smaller or larger than this. Rather than comparing this attribute across the whole comparative collection, it is more useful to consider each of the typologies for bundling morphology. When the range of specimens containing each type of bundling arrangement is compared with one another, bundle length becomes an important attribute in differentiating between them.

Within the atactostele collateral typology Dioscorea alata has statistically larger bundles than the other five specimens. Where the other specimens have bundle lengths that range between approximately 150-550µm, D. alata has a range from 250-890µm. On the opposite end of the spectrum, Saccharum officinarum is the only specimen with this bundle typology that also has bundles smaller than 150µm. It is harder to differentiate between the specimens with atactostele amphivasal arrangement. There is overlap between the bundle length ranges of all specimens apart from Xanthosoma sagittifolium which has the bundles that can be smaller than 150µm, with a minimum size of 100µm. Both Zingiber spp. within the atactostele amphicribal bundle typology have statistically different bundle length ranges. Zingiber sp. (BG957) has a range from 300-450µm, while Zingiber sp. (EU008) has a range smaller than this between 160-300µm.

The fruit specimens can also be differentiated based on the ranges of bundle lengths within the bundle typologies. The specimens with amphivasal bundles have some overlap between 210-750µm, but the box plot shows that there some specimens with smaller bundles that can be differentiated from the others. Barringtonia asiatica has very little overlap with any other species, with a range of 85-160µm which represents the smallest bundles in this typological category. Slightly larger than this is Barringtonia racemosa, which has a range of 150-410µm and therefore can be identified based on bundle lengths from 160-210µm. The remaining Pandanus spp. specimens within this bundle typology cannot be differentiated based on bundle length. Amphicribal specimens have a much larger amount of overlap between the bundle length ranges of specimens. At the smaller end of the spectrum, bundles smaller than 170µm can belong to either Artocarpus altilis or Musa sp. 1. Bundles larger than 620µm can only belong to Musa sp. 2 according to these statistical ranges.

Figure 5.14 Box plot showing vascular bundle lengths within reference collection according to tissue arrangement

Combining these two morphological attributes allows more accurate differentiation of parenchyma in the comparative collection. For those specimens that had similar bundle lengths to others with the same bundle typology, a multivariate approach was needed to further test whether classification of unidentified samples is possible based on bundle morphology. This multivariate statistical approach included an additional variable- bundle width- within a PAST dataset and used the same DFA approach as that used to explore starch and parenchyma cell morphology. First, individual datasets were created for the atactostele (stem-derived roots and tubers) and the fruits so that these could be analysed separately. Two ordinal variables were included- bundle length and width, and one nominal- bundle morphology.

The specimens within the atactostele bundle typology dataset had an overall correct reclassification rate of 74.7%. A number of specimens had all five recorded bundles correctly re-classified into the same specimen. These included Araceae (EU2012-08), Cordyline fruticosa, Dioscorea esculenta, Saccharum officinarum, Tacca leontopetaloides, Xanthosoma sagittifolium and Zingiber sp. Several others only had one bundle incorrectly classified, such as Dioscorea nummularia, Epipremnum pinnatum and another Zingiber sp. Alocasia macrorrhiza, Cyrtosperma merkusii, Dioscorea alata and X. sagittifolium all had 60% (n=3) of the five bundles reclassified correctly, while the remaining two specimens had less than half of the bundles correctly assigned and were not easily discriminated from the other specimens based on 94 these three attributes. The overall loadings from this analysis indicated that bundle length was consistently the most diagnostic variable included in the dataset.

The confusion matrix constructed indicated that the specimens in fruit bundle typologies were less able to be distinguished from one another based on bundle length, width and arrangement, than the atactostele specimens. Overall, 70.2% of the bundles included in this dataset were correctly reclassified to specimen of origin. As with the atactostele dataset, there were a number of specimens that had all of the included bundles correctly re-assigned. These specimens included Artocarpus altilis, Barringtonia asiatica, Musa sp.1, and Syzygium malaccense. Several others had over half of the bundles correctly reclassified to original species, including Musa sp. 2 and Pandanus tectorius with 80% correct. The outputs of using DFA on this dataset indicate that these particular specimens are able to be differentiated from the others within the dataset based on the three attributes for bundle morphology used here. Bundle length was again shown to be the most diagnostic variable within the loadings for the fruit dataset.

Consideration of the diagnostic value of these attributes within the atactostele and fruit datasets separately is useful for discrimination when the arrangement of vascular bundles is known; however when attempting to identify a fragment of archaeological parenchyma, the overall arrangement is not always visible within that particular fragment. Considering this, the value of bundle morphology in the dataset as a whole was assessed. The atactostele and fruit datasets were combined and the DFA run again with the same three variables of bundle length, width and arrangement. Overall, the percentage of correct re-classifications within this dataset was 62.3%. Eight specimens had all five recorded bundles correctly re-assigned, including Artocarpus altilis, Araceae sp., Barringtonia asiatica, Dioscorea esculenta, Musa sp. Syzygium malaccense, Saccharum officinarum, and Tacca leontopetaloides. These were all specimens that had 100% correct re-classifications within the separated datasets and so are easier to classify when compared to the whole range of specimens with vascular bundling in the comparative collection. Another five specimens had four out of the five bundles re-classified to the correct specimen of origin. These included Cordyline fruticosa, Epipremnum pinnatum, Musa sp.2, Pandanus tectorius and Zingiber sp. These and a further five specimens, Alocasia macrorrhiza, Cyrtosperma merkusii, Xanthosoma sagittifolium, Zingiber sp. had over half of the included vascular bundles allocated to the original specimen and could therefore be identified based on these three attributes with a moderate level of confidence. The remaining seven species could not be confidently differentiated from the other specimens included in the analysis. These specimens include Barringtonia racemosa, Colocasia esculenta, Dioscorea alata, Dioscorea nummularia, Ficus copiosa, Pandanus tectorius and Tabernaemontana aurantiaca.

Cell Cell Vascular length width Vascular bundle Species Organ Boundary tissue description Ground tissue description bundle length Vascular tissue description range range width range (µm) range (µm) (µm) (µm) Outer epidermis of 2-3 layers of thickened Rounded and isodiametric, angular and Artocarpus altilis Fruit radially organised rounded and isodiametric 40-60 30-40 elongate, rounded and elongate. Inter- 100-600 70-280 Atactostele and amphicribal u-shaped. cells. cellular spaces. Outer thin epidermis composed of 1 layer of Mostly rounded and isodiametric, some Artocarpus altilis Seed radially organised rounded and isodiametric 60-80 50-65 elongate. Inter-cellular spaces. Pith of 80-900 75-360 Atactostele and amphicribal concentric. cells with thickened cell walls. sclerenchyma. A layer of thick periderm. Cortex of c.20 rows of angular and elongated cells with very thin Mostly rounded and isodiametric, some Alocasia macrorrhiza Corm 80-130 70-100 300-560 200-450 Atactostele and amphicribal open-ended. cell walls. Vascular cambium of 2 rows of elongate. Inter-cellular spaces. similar cells with thicker walls. Outer epidermis of 3 layers of radially Rounded and elongate, some Angiopteris sp. Rhizome organised rounded and elongated cells with 100-150 85-100 Unmeasured Unmeasured Dictyostele. isodiametric. Inter-cellular spaces. thickened cell walls. Outer epidermis of one layers of radially organised angular and isodiametric cells with Rounded and isodiametric, some Asplenium sp. 1 Rhizome 55-80 45-70 770-960 390-810 Dictyostele and amphicribal concentric. thickened cell walls. Below this a broad region elongate. Inter-cellular spaces. of 7-8 rows of fibres. Outer epidermis of one layers of radially organised angular and isodiametric cells with Angular and isodiametric or elongate. Asplenium sp. 2 Rhizome 115-175 90-130 160-950 110-570 Dictyostele and amphicribal concentric. thickened cell walls. Below this a broad region Inter-cellular spaces. of 7-8 rows of fibres. Outer epidermis of one layer of very small radially organised rounded and isodiametric Angular and isodiametric. Inter-cellular Barringtonia asiatica Fruit 55-105 40-75 85-160 85-150 Eustele and amphivasal concentric. cells with a thick outer cell wall. Below this a spaces. row of palisade mesophyll cells. Outer epidermis of one layers of very small radially organised rounded and isodiametric Rounded and isodiametric, some Barringtonia asiatica Seed cells with thick cell walls. Two rows of 50-70 35-50 Not observed Not observed Not observed elongate. Inter-cellular spaces vascular cambium separates the cortex and pith. Thin epidermis of a single layer of angular and elongated cells with thick cell walls. Rounded and isodiametric, some Barringtonia racemosa Fruit Another possible 10 rows of rounded and 45-65 35-50 150-400 110-300 Eustele and amphivasal concentric. elongate. Inter-cellular spaces. elongated cortical parenchyma below the epidermis. Outer epidermis of one layers of rounded and isodiametric cells with a thick outer cell wall. Angular and isodiametric. Inter-cellular Barringtonia racemosa Seed Two rows of very small angular and elongated 60-85 50-70 Not observed Not observed Not observed spaces. cells make up the vascular cambium which separates the cortex and pith.

Outer thickened epidermis composed of 3 layers. 20 rows of cortical parenchyma angular and elongated with thin cell walls. Rounded or angular and isodiametric or Atactostele and amphivasal concentric or Colocasia esculenta Corm 70-100 50-75 230-580 160-400 Two rows of angular and isodiametric cells elongate. Inter-cellular spaces. u-shaped. with thick cell walls make up the vascular cambium.

Table 5.9 Summary of parenchyma morphology within reference collection 96

Outer thickened epidermis and below this a Angular and isodiametric or elongate. Atactostele and amphivasal concentric. Cordyline fruticosa Root region of cortical parenchyma which is 55-75 40-55 200-300 100-200 Inter-cellular spaces. Fibre sheath. rounded and isodiametric with thin cell walls. Rounded and elongate, some Cyrtosperma merkusii Corm Not observed 40-80 30-50 300-500 200-300µm Atactostele and amphivasal open-ended. isodiametric. Inter-cellular spaces. Rounded and elongate or isodiametric, Epidermis consisting of 4 layers of angular Dioscorea alata Root tuber 80-110 50-75 some irregularly rounded and elongate. 230-880 85-340 Atactostele and closed collateral. and elongated cells with thick cell walls. Inter-cellular spaces. One layer of thickened epidermis. A region of Aerial Rounded and isodiametric, some Dioscorea bulbifera vascular cambium of five rows of angular and 95-130 75-100 Not observed Not observed Not observed bulbil elongate. elongated cells. Epidermal region of 3 layers of angular and Rounded and isodiametric, some Dioscorea esculenta Root tuber 150-235 100-160 150-380 80-240 Atactostele and closed collateral. elongated cells. elongate. Some inter-cellular spaces. Thickened epidermis outside 5 rows of Rounded and isodiametric, some Dioscorea nummularia Root tuber 75-115 60-100 155-435 100-250 Atactostele and open or closed collateral. angular and elongated cortical parenchyma. elongate. Inter-cellular spaces. Thickened single-layer periderm above 5 rows of angular and elongated primary cortical parenchyma cells with thin cell walls. Another Rounded and isodiametric, some Epipremnum pinnatum Corm five rows of similarly shaped cells compose a 70-95 50-70 220-375 180-300 Eustele and amphivasal u-shaped. elongate. Inter-cellular spaces. region of secondary cortex. A row of vascular cambium of one layer of rounded and elongated cells. Rounded and isodiametric. Inter- Eustele and closed collateral or Ficus copiosa Fruit Not observed 40-60 30-45 70-380 60-150 cellular spaces. amphicribal u-shaped or concentric. Angular and isodiametric or elongate. Ficus tinctorius Fruit Not observed 55-90 25-70 Not observed Not observed Not observed Inter-cellular spaces. The parenchyma within the secondary xylem same as phloem, regions of A region of periderm of about 4 rows of Region of parenchymous secondary anomalous tertiary growth adjacent to Ipomoea batatas Root tuber radially-orientated angular and isodiametric 80-115 60-90 phloem outside the cambium. Angular Not applicable Not applicable individual vessels. Divide both cells with thin cell walls. and isodiametric, some elongate. periclinally and tangentially to produce concentric radiating rings of tertiary xylem. A layer of endocarp consists of a row of Rounded and isodiametric, some Inocarpus fagifer Seed angular and isodiametric cells with very thick 45-100 35-75 Unmeasured Unmeasured Atactostele and amphicribal u-shaped. elongate. cell walls. Rounded and isodiametric, some Morinda citrifolia Fruit Not observed 40-60 30-50 Not applicable Not applicable Not applicable elongate. Irregularly rounded and isodiametric, Musa sp. 1 Fruit Not observed 60-90 50-70 some elongate. Inter-cellular spaces and 80-115 70-100 Eustele and amphicribal concentric. duct cavities. Irregularly rounded and isodiametric, Musa sp. 2 Fruit Not observed 73-215 46-170 some elongate. Inter-cellular spaces, 220-680 180-370 Eustele and amphicribal concentric. duct cavities and fibre bundles.

Cell Cell Vascular length width Vascular bundle Species Organ Boundary tissue description Ground tissue description bundle length Vascular tissue description range range width range (µm) range (µm) (µm) (µm) Epidermis of one row of thickened, rounded Fruit and isodiametric cells. Below is one row of Rounded and isodiametric, some Atactostele and amphicribal concentric. Pandanus tectorius 60-100 40-70 260-750 210-430 phalange similarly rounded and isodiametric cells with elongate. Inter-cellular spaces. Concentrated near pith. Fibre sheath. slightly thinner cell walls. Periderm made up of around 6 rows of angular and elongated cells. Below is a region Rounded and isodiametric, some Xylem vessels are present near the pith of Pangium edule Fruit 40-80 30-60 Not applicable Not applicable of parenchyma interspersed with elongate. Inter-cellular spaces. the fruit. sclerenchyma. Secondary xylem same as those within the Region of periderm of about 5 rows of angular phloem, however there are regions of and isodiametric cells with thin cell walls. The anomalous tertiary growth adjacent to Primary Rounded and isodiametric, some Pueraria lobata cambium separates the cortex from the stele, 25-45 20-30 Not applicable Not applicable individual vessels. Polyarch xylem. Many root elongate. Inter-cellular spaces. and is composed of an endodermis and areas of tertiary xylem and phloem are pericyle of one row. also present and are dissected by medullary rays. Primary tissues made up of angular and isodiametric cells with very few inter- cellular spaces. Within this, wide medullary rays made up of ligneous fibres and also xylem vessels. Scanty paratrachial to vasicentric layers of Primary Region of periderm of 3 rows of angular and Rounded and isodiametric. Inter- Piper methysticum 45-75 35-60 Not applicable Not applicable xylem cells abut these rays and are root isodiametric cells that are thick walled. cellular spaces (pith). differentiated by thinner walls and more angular shape. Bundles of phloem are contained within these areas of xylem. Exterior to the cambium are further bundles of xylem within the parenchymous tissues. Outer epidermis of around 3 layers of rounded Angular and elongate, some Pteridium sp. Rhizome 80-115 65-90 Unmeasured Unmeasured Dictyostele. and elongated cells with thickened cell walls. isodiametric.

Outer thickened epidermis, cortex consists of about 4-5 rows of angular and elongate cells Irregularly rounded and elongate or Solanum tuberosum Stem tuber 90-130 70-100 Unmeasured Unmeasured Atactostele and bicollateral. with thinner walls. The vascular cambium is isodiametric. two layers of angular cells. Outer thickened periderm, and a cortical Angular and elongate, some Eustele and closed collateral or Spondias dulcis Fruit region of approximately 5-10 rows of rounded 85-160 55-95 280-400 140-240 isodiametric. Thin cell walls. amphicribal concentric. and isodiametric cells with thick cell walls.

Thin epidermis of a single row of angular and elongated cells, with a thicker exterior wall. Angular and elongate, some Eustele and amphicribal stellate with Syzygium malaccense Fruit Several rows of small cells that are angular in 75-120 45-85 250-620 150-330 isodiametric. Few inter-cellular spaces. polyarch xylem inside bundle. shape and more isodiametric in dimension are below this within the cortex Tabernaemontana Epidermis a single row of tangentially Rounded and elongate or isodiametric. Fruit 35-60 25-40 175-370 100-230 Eustele and amphicribal concentric. aurantiaca flattened angular and elongated cells Inter-cellular spaces. 98

Cell Cell Vascular length width Vascular bundle Species Organ Boundary tissue description Ground tissue description bundle length Vascular tissue description range range width range (µm) range (µm) (µm) (µm) Periderm of 3 rows of r angular and elongated Rounded and isodiametric, some Tacca leontopetaloides Stem tuber cells with thin cell walls, with the outermost 70-100 60-80 250-530 100-270 Atactostele and bicollateral. elongate. Inter-cellular spaces row having a thicker external wall Outer epidermis of one layers of angular and Angular and isodiametric, some Todea sp. Rhizome isodiametric cells with thickened cell walls. 75-135 60-105 Unmeasured Unmeasured Dictyostele. elongate. Below is a broad region of fibres. Periderm of 4 rows of angular and Xanthosoma isodiametric cells with thin cell walls, with Rounded and isodiametric. Inter- Atactostele and amphivasal concentric. Corm 55-80 50-60 90-270 80-230 sagittifolium the outermost row having a thicker external cellular spaces. Fibre sheath. wall. Thickened periderm with around 10 rows of Primary Irregularly rounded and isodiametric or Eustele and closed collateral. Fibre Zingiberaceae sp. 1 thick walled cells that are rounded and 55-80 50-60 296-447 240-352 root elongate. Inter-cellular spaces. sheath. broadly isodiametric in dimension. Periderm of around 10 rows of rounded and elongated thick walled cells. Thicker exterior Primary Rounded and isodiametric, some Eustele and amphicribal concentric. Fibre Zingiberaceae sp. 2 wall of the outermost layer. Vascular 75-105 50-85 160-309 112-240 root elongate. sheath. cambium that is a single layer of more angular and elongated cells.

Description of charcoal Each of the specimens in the modern comparative collection was experimentally charred from both dried and fresh state to assess the nature of morphological changes in the boundary, ground and vascular tissues. SEM images provided high resolution records of the results of the charring process. These morphological changes were described at an individual level for each specimen in Appendix A, and are summarised briefly here. The conditions of charring were described earlier within this chapter (see section ‘Experimental charring’), but it is important to note that the temperature, time and access to oxygen were kept consistent for all of the specimens. The charred specimens represent only one experiment in charring conditions, and morphological changes may be different under alternative charring conditions of different temperatures, length of exposure to heat, and oxidising conditions.

Boundary tissues A small number of changes can occur in boundary tissues during charring. The epidermis or periderm often becomes compressed or alternatively transforms into a region of solid carbon. The carbonisation of the boundary tissues occurs in stem and root-derived tissues, as well as fruits primarily within the samples charred from fresh state. There are also examples of this carbonisation occurring in the samples that were dried prior to charring such as that within Asplenium sp., Cordyline fruticosa, Dioscorea bulbifera, Inocarpus fagifer, Morinda citrifolia, Spondias dulcis and Tabernaemontana aurantiaca. The remaining samples charred from a dried state had epidermal or peridermal tissues that were either compressed or retained their original morphology. Similarly, those tissues within samples charred from a fresh state were either turned to solid carbon or were observed to be in the same condition as the uncharred sample. These observations indicate that the samples charred from a dry state were more consistent in terms of morphological changes than those charred from fresh state.

Ground tissues Ground or conjunctive tissues can undergo a range of different morphological changes when charred from fresh and dried states. First, the shape of the cells can change. Seven specimens in the comparative collection evidenced cell shape change. One specimen, Barringtonia asiatica, had more rounded cells rather than angular when dry-charred. The cells of another specimen, Asplenium sp. became more angular in both samples. The remaining specimens had either partly or wholly cells in the ground tissue that were originally rounded and had become more irregularly rounded. These included Alocasia macrorrhiza, Dioscorea alata, Dioscorea nummularia, Pandanus tectorius and Zingiber sp. in the dry-charred samples, but only A. macrorrhiza, and another sample of Pandanus tectorius and Zingiber sp. in the wet-charred samples.

Many specimens also had either inter-cellular spaces or whole regions of ground tissue transform to solid carbon. This change was only slightly more common in the wet-charred 100 samples, which is an unexpected observation. The wet samples are more hydrated when exposed to the high temperature during charring, and so greater modification is usually observed in the form of carbonisation and tension fractures as the liquid evaporates. Carbonisation of inter-cellular spaces was observed within six dry and wet-charred specimens, but these were not the same specimens in both instances. Within the dry-charred samples, these included Alocasia macrorrhiza, Artocarpus altilis seed, Barringtonia racemosa seed, Dioscorea bulbifera, Ficus copiosa and Musa sp.2. The specimens with this variety of carbonisation in the wet-charred samples included A. macrorrhiza, Colocasia esculenta, Dioscorea alata, .D. bulbifera, F. copiosa and Musa sp.2. Almost half of the modern reference collection also had regions or complete carbonisation of the ground tissue. Of the total 36 specimens, 15 specimens had carbonisation in the dry-charred samples (42%), and 16 specimens had carbonisation in the wet- charred samples (44%).

Cavities, tension fractures and vesicularisation also often occurred within both the wet and dry- charred samples from the evaporation of water within the samples. The process of cavity formation and fracturing is called rhexigeny (Hather 2000:41, 1993:4) and involves the mechanical tearing of tissues under tension. The location of these tears in the ground tissue can vary. Some were observed within the cambium, separating the cortex from the pith, while others were near vascular tissues or were randomly located throughout the pith. Cavities formed within 12 of the dry-charred specimens, but were much more common in the wet-charred samples. Over half of the specimens in the comparative collection had cavities form in the wet-charred samples. This modification was observed in a total of 18 specimens. These included A. altilis seed, Angiopteris sp., Barringtonia asiatica seed, B. racemosa seed, C. esculenta, Musa sp 1.and Xanthosoma sagittifolium, which only form cavities when charred from fresh state. Alternately, Cyrtosperma merkusii only formed cavities within the dry-charred sample. Tension fractures were also very common, forming in eight dry and wet-charred samples, but again these were not the same specimens in both types of samples. Only Musa sp. 2., Pueraria lobata, Pteridium sp., Spondias dulcis and X. sagittifolium had cavities form in both wet and dry- charred samples.

Vesicularisation is the formation of air spaces by the explosion of cells under pressure. Hather (2000:45) describes this process as occurring when “...the water content of cells heats up and eventually boils, resulting in the rupture of the cell and a release of water vapour under pressure. The escaping water vapour bubbles through the tissue, compressing cells and causing the formation of vesicles.” The vesicular appearance is a by-product of the fact that some of the cells are still visible as such but become much smaller and the cell walls are smooth with the outline of cells on the internal surface. This form of cellular modification was observed in nine dry-charred specimens and eight dry-charred specimens. These were mostly the same specimens in both, with the exception of Angiopteris sp., Colocasia esculenta, and Cyrtosperma merkusii 101 which were only vesicular in the dry-charred samples and Dioscorea nummularia, which only had vesicles in the wet-charred samples.

A number of changes can also directly occur within the ground tissue cells and affect the appearance of these cells. Roughly one third of the reference collection had samples that exhibited cell compression either regionally or throughout the ground tissue in the dry-charred samples (n=12), and half of the collection had this modification within the wet-charred samples. This was usually a by-product of the formation of cavities and tension fractures within these tissues. Those specimens that only had cell compression within the wet-charred samples included Angiopteris sp., Asplenium sp., C. esculenta, Ipomoea batatas, Musa sp.1, Tacca leontopetaloides, and Todea sp. A smaller number of samples only had compression in the samples charred from dried state, including Dioscorea esculenta, Inocarpus fagifer, Pueraria lobata and Xanthosoma sagittifolium.

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Cells Cells Cells Inter-cellular Tension Compression of Thicker cell Thinner cell Fractured/collapsed Species rounded angular irregular spaces carbonise Cavities fractures cells walls walls Vesicularisation Carbonisation cells Shallower cells Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Alocasia macrorrhiza X X X X X X X X Artocarpus altilis fruit X Artocarpus altilis seed X X X Angiopteris sp. X X X X X Asplenium sp. 1 X X X X X Asplenium sp. 2 X X X X Barringtonia asiatica seed X X X Barringtonia racemosa seed X X X X X Colocasia esculenta X X X X X Cordyline fruticosa X X X X X X Cyrtosperma merkusii X X Dioscorea alata X X X X X Dioscorea bulbifera X X X X X X X X X X Dioscorea esculenta X X X X Dioscorea nummularia X X X Epipremnum pinnatum X X X X Ficus copiosa X X X X X Ficus tinctorius X X X X Ipomoea batatas X X X X Inocarpus fagifer X X X X X X Morinda citrifolia X X x X X X Musa sp. 1 X X X X X Musa sp. 2 X X X X X X Pandanus tectorius X X X X Pangium edule X X Pueraria lobata X X X X Piper methysticum X X Pteridium sp. X X X X X X Spondias dulcis X X X X X X X X Syzygium malaccense X X X X X X X X Tabernaemontana aurantiaca X X X X X X Tacca leontopetaloides X X X X X Todea sp. X X Xanthosoma sagittifolium X X X X X X X Zingiberaceae sp. 1 X X X X X X Zingiberaceae sp. 2 X X X X X X

Table 5.10 Description of morphological modification within ground tissue of charred samples in the comparative collection 103

Phloem- cavity Phloem- carbon Bundle separates Cambium- carbon Fibre sheath-carbon Phloem compressed Cambium fracture Species Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Dried Fresh Alocasia macrorrhiza X Artocarpus altilis fruit X X Artocarpus altilis seed X X Angiopteris sp. X X Asplenium sp. 1 X X X Asplenium sp. 2 X X X X Barringtonia asiatica seed Barringtonia racemosa seed Colocasia esculenta X X Cordyline fruticosa X X X Cyrtosperma merkusii X X X Dioscorea alata X X Dioscorea bulbifera Dioscorea esculenta X X X Dioscorea nummularia X X Epipremnum pinnatum X X Ficus copiosa X X Ficus tinctorius Ipomoea batatas Inocarpus fagifer X X Morinda citrifolia X X Musa sp. 1 X X Musa sp. 2 X X Pandanus tectorius X X Pangium edule X X Pueraria lobata X X X Piper methysticum Pteridium sp. Spondias dulcis X X Syzygium malaccense X X Tabernaemontana aurantiaca Tacca leontopetaloides X X Todea sp. X X Xanthosoma sagittifolium Zingiberaceae sp. 1 X X X Zingiberaceae sp. 2 X X

Table 5.11 Description of morphological modification within vascular tissue of charred samples in the comparative collection

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Cells were also often fractured or collapsed in many of the wet and dry-charred samples. This process often rendered regions of the ground tissue unrecognisable. Many of the fruits exhibited this change, including Ficus tinctoria, Morinda citrifolia, Musa sp. 1, Pangium edule, Pandanus tectorius, Spondias dulcis, Syzygium malaccense and Tabernaemontana aurantiaca within either the wet or dry-charred samples. A small number of stem and root- derived specimens had large areas of collapsed cells with exposed cell contents such as starch, crystals and druses. These included Cordyline fruticosa, Epipremnum pinnatum, Ipomoea batatas and Zingiber sp. In addition, a very small number of specimens exhibited signs that cells became shallower during dry-charring, indicating a change in cell dimension. These included Dioscorea esculenta and I. batatas.

Where specimens did not show signs of cell compression or collapse, many instead had thickening or thinning of the cell walls. This can result from cells swelling and solidifying during intense heating of liquids within the tissues. Within the dry-charred samples, Alocasia macrorrhiza, Artocarpus altilis seed, Barringtonia asiatica seed, Colocasia esculenta, and Dioscorea bulbifera all had cell walls that were thicker than those observed within the histological thin sections. Only A. macrorrhiza and D. bulbifera had this occur within the samples charred from fresh state. Conversely, a small number of samples had thinning of the cell walls after charring. These included D. alata and Zingiber sp. in the dry-charred samples, and D. esculenta and Zingiber sp. within the wet-charred samples. The sample of C. fruticosa that was charred from dried state did not show signs of thickening or thinning of cell walls, but instead had a pitted texture on the interior of the cell lining.

Vascular tissues A smaller array of changes occurred in the vascular tissues during the charring process. The root-derived specimens experienced less modification in parenchyma morphology. Only Pueraria lobata had some rhexigenous tension fractures and carbonisation occur within the vascular cambium, separating the secondary xylem from the phloem. This occurred in both the wet and dry-charred samples of the species. The vascular bundles of the stem-derived specimens and fruits often had significant morphological changes occur through charring. The thick non- living lignified walls of the xylem are able to withstand the intense heat of the charring process much better than the more fragile thin living tissues of the phloem that often contains sugars and nutrients. The xylem was therefore consistently preserved in original condition, but the phloem either turned to solid carbon or became a cavity.

The phloem became a cavity within 11 of the dry-charred specimens and 12 of the wet- charred specimens, which represents just over a third of the 33 fruits and stem-derived charred specimens. These included most of the yams (Dioscorea spp.) including Tacca leontopetaloides and both Musa spp. Another 14 specimens exhibited the fusion of tissues within the phloem in the dry-charred samples, consequently transforming these tissues into solid carbon. These 105 included both the fruit and seed of Artocarpus altilis, several of the Pteriodophytes, and three of the aroids. Colocasia esculenta, Cyrtosperma merkusii and Xanthosoma sagittifolium had this change occur within the phloem, but interestingly the only other aroid in the comparative collection, Alocasia macrorrhiza, did not exhibit any changes in the dry-charred samples made from the corm. However, change did occur within the wet-charred sample of A. macrorrhiza. It is also interesting to note that the phloem within the corm of C. merkusii did not exclusively turn to solid carbon in the dry-charred samples; some regions of the phloem were instead compressed. This could indicate differential charring within the sample. Some specimens such as Dioscorea bulbifera did not have recognisable vascular bundles within the charred samples (this could be result of the sample selection and fracturing process) and thus could not be described here. It is important to note that the arrangement of vascular tissues within the root and stem-derived tissues and fruits did not change under charring. The xylem is still identifiable as such, and the presence of the phloem as either carbon or a cavity enables description of the arrangement of vascular tissues, and also the organisation of these tissues within the organ where the fragment is large enough.

Other changes that could occur within the vascular bundles included the formation of a cavity abutting the bundles, where these tissues have broken away from the surrounding conjunctive tissue. This was only noted in a small number of specimens of the dry-charred samples, which included Cordyline fruticosa, C. merkusii and Dioscorea esculenta. The fibre sheaths surrounding the vascular bundles of the rhizomes of Asplenium spp. and Zingiber sp. also turned to solid carbon within the dry-charred samples. Only one Asplenium species exhibited this variation of tissue carbonisation within the wet-charred samples.

In summary, the morphological changes that can occur during charring need to be considered during the identification process. Some significant changes to cell and tissue morphology can occur when plant specimens are charred either from dried or fresh states and these changes will inhibit any identification that is based solely on morphological patterning in fresh samples. However, these changes described here are based upon particular charring conditions, and so it is hard to gauge how these might differ under alternative conditions such as different temperatures, length of exposure and oxidising conditions. An awareness of the nature of these changes and an understanding of why these might occur are essential components of a methodology that will aid classification of unknown archaeological parenchyma to a taxonomic level with confidence.

Development of an Identification Flowchart Key Through exploration of the morphology of ground tissues and vascular tissues separately, it becomes clear that many taxa in the comparative collection are able to be differentiated from one another. These tissues are the components of parenchyma most likely to be observed and

106 preserved within the archaeological record, and therefore the attributes analysed above can aid identification of these specimens with a moderate-to-high level of confidence. Where taxa could not be differentiated based on the morphology of one of these tissues, an identification flowchart key is useful that combines all of the known diagnostic features of parenchyma within the reference collection. Depending upon the presence or absence of vascular tissues, the flowchart can guide identification of an unknown sample to various taxonomic levels based firstly upon the arrangement of vascular tissues, then bundle morphology, and finally considering cell morphology. If one attribute cannot be observed, another avenue allows identification using other attributes. Sometimes it is only possible to narrow the list of potential classifications, and the key is the first step in identifying unknown parenchyma. Further examination of SEM and light microscopic imagery, along with consideration of geographic and environmental boundaries for plant distribution can aid final identification.

Essentially, two flowcharts were created based on the morphological data collected from the thin sections made from each specimen in the comparative collection. The first key is the primary flowchart, and is usable if some vascular tissues are visible within an unidentified fragment of desiccated or charred parenchyma. This chart starts by considering the arrangement of vascular tissues, then the morphology of bundling, and finally the cell morphology is used to differentiate between taxa. If particular attributes are not able to be observed, the key can at least facilitate classification to the type of organ from which it derived (fruit, stem-derived or root-derived), or to family level in some cases. There is more morphological patterning within the vascular tissues of stem-derived tissues at family level, and particular vascular arrangements can be seen throughout the specimens of a family. For example, collateral vascular bundles are present in the members of the Dioscoreaceae family within the comparative collection, while amphivasal bundles are present in all members of the Araceae family. The specimens of fruits included in the comparative collection also have these types of bundles, but other aspects of cellular morphology and charring characteristics differentiate these at family level.

The second key is usable if vascular tissues cannot be observed within the sample and only uses attributes of cell morphology such as shape and cell lengths. Attributes such as the presence of fibres, vessels, duct cavities and inter-cellular spaces were also used when possible. Clearly some changes to boundary, ground and vascular tissues occur during charring and those observed changes were incorporated into this key. This key may only be able to provide a list of tentative classifications that are not usually from the same genus or family. This is because there is a lot less morphological patterning within the ground tissue at these taxonomic levels. There is significant overlap between fresh specimens, but particular characteristics of charring can be used to differentiate these to some degree.

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Sometimes several directions within the flow charts could be taken to arrive at the same classification. This is because some species have several cell shapes or bundle arrangements, or charring could affect cell shape or dimensions. If a junction is reached where the particular attributes to further narrow down the list of possible classifications are not visible, then the data in Appendix A containing detailed descriptions of the fresh and charred samples can be examined. Where boundary tissues are present, these descriptions can be used instead to differentiate between this list. Photographs and SEM images of the reference collection then needed to be considered along with geographical data for the distribution of each of the species. It was deemed useful here to adopt the determination system using identification criteria and associated levels of confidence compiled by Paz (2001) and later used by Oliveira (2008). This determination system has been used successfully by these researchers in the past to analyse macrobotanical plant remains from a range of archaeological contexts in the Asia-Pacific region (Barker et al. 2011; Barton and Paz 2007; Oliveira 2008; Paz 2001, 2005). Plant samples are determined to taxa using the following criteria:

Non prefixed: Photographic reference(s) and/or illustration reference(s); reference material not essential; exact fit of the taxonomic features, geographic distribution, and species citation in the local flora;

Prob.: Flora citation, geographic area compatibility, an agreement with taxonomic details; image OR illustration OR reference material (not necessarily an exact or good fit);

Cf.: All six categories may or may not exist; archaeological specimen resembles image OR illustration OR reference material OR previous identification; flora, taxonomic details and geographic area but with doubts;

Elim.: Lowest level of confidence for a binominal determination to species level, but with no access to image, illustration or reference material; taxonomic description, geographic area and other species of same genus were eliminated from local/regional flora (= likely candidate);

Suffix 'type': Very low level of confidence, used only at family and genus level of determination; shape of specimen fits the geographic distribution, some morphological characters, and may be in the local flora;

Form shape description: None of the six types of information exist (image, illustration, reference collection, flora, taxonomic details and geographic area), but the specimen is distinctly a seed, nut fragment or a certain plant part.

The identification process for archaeological parenchyma incorporated a large amount of data derived from the modern comparative collection to provide classifications with varying levels of confidence, expressed using the determination system of Paz (2001). The two

108 identification flowchart keys enable the classification of unknown plant samples that have varying levels of tissue preservation to at least the type of organ from which fragments have derived. This information is crucial to the interpretation of plant use within a site or landscape. The presence of crops is a good indicator of at least low-level agricultural practices. Here, this information will be used as stand-alone data and to corroborate the data collected from the analysis of soil samples for ancient starch residues. The presence of starch and parenchyma from a particular family, genus or species will provide complementary records for the presence of economic and famine species within the sites of Talasiu, Leka and Heketa on Tongatapu in the Kingdom of Tonga.

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Figure 5.15 Flowchart 1 used as an identification key to identify unknown parenchymatous samples when vascular tissues are visible 110

Figure 5.16 Flowchart 2 used as an identification key to identify unknown parenchymatous samples when no vascular tissues are visible 111

PART TWO- DEVELOPMENT OF PREHISTORIC AGRICULTURE IN TONGA

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Chapter 6 Sites and Field Sampling Strategy

This chapter outlines the methods used to extract botanical remains from three archaeological sites on Tongatapu of varying antiquity. The archaeobotanical program builds upon the work carried out to date within Polynesia (Chapter 2) and attempts to answer the key research questions regarding the role of plants within colonising subsistence and migration episodes in Western Polynesia and the later development of social complexity through the establishment of the maritime Tu’i Tonga chiefdom. Revised protocols are established for both field and laboratory methods, building on those currently used both in the Pacific and elsewhere (Chapters 3-4) through new experimentation. Experiments were conducted in reaction to problems highlighted within previous studies in terms of the success of extractions within differing deposits, and the risk of contamination from laboratory equipment. The results of these experiments were used to provide data on the presence of plant taxa within well-dated cultural deposits that are presented and discussed in Chapters 8 and 9.

Methodology for field sampling of archaeological sediments

Site selection During fieldwork in November 2011, sediments were sampled from open archaeological sites using flotation and bulk sampling for microbotanical analyses. Comparisons of identified taxa within sediments were made on a spatial scale across north-eastern Tongatapu, to provide a scope for the chronology of the introduction of crops and development of production systems. This incorporated test-pitting at two known sites on the south-eastern edge of the Fanga’Uta Lagoon, and one site at Heketa on the northern coast underneath the ‘esi or chiefly back-rest monolith. Talasiu (TO-Mu-2) and Leka (J17) are both located in the Mua region within the village of Lapaha and are now inland although Talasiu is located on the edge of an old palaeoshore (Dickinson 2007). The ‘esi (TO-Nt-2) is attributed to the construction phase at Heketa initiated by the 11th Tui Tonga within local traditions, and is considered contemporary with the Ha’amonga trilithon of the same complex.

These sites had already been located, surveyed and sampled for archaeological material by Clark and others (2008), Golson (1957), McKern (1929), Poulsen (1987), and Spenneman (1986, 1989) in archaeological projects on Tongatapu. McKern (1929) carried out the first major archaeological survey of the Tongan archipelago as part of the Bayard Dominick Expedition of the Bernice P. Bishop Museum in 1920-21. Mapping and excavation of the tombs and enclosing ditch at Lapaha and Heketa was a particular focus of his work, and his labelling of the tombs has been used in all subsequent archaeological research. Langi Leka (J17) and the ‘esi at Heketa (TO-Nt-2) were among the monuments mapped by McKern. Golson (1957) carried out a limited survey of Tongatapu and offshore islets, exploring the potential for using ceramics as cultural markers. During this survey Golson excavated a section of a site named by 113

Spenneman TO-Mu-2 near the village of Talasiu, and reported on the potential of the pottery- bearing midden. Golson encouraged one of his students, Jens Poulsen, to further explore the ceramic typological sequence within Tongatapu. Poulsen (1964, 1967, 1987) subsequently spent a year carrying out a detailed program of excavation around the Fanga’Uta lagoon.

Following extensive archaeological research in the late 1960s by Davidson (1969, 1971), Groube (1971) and others that focused on late prehistoric sites and the construction of a Tongan culture history, Spenneman (1986) spent some months in Tonga attempting to locate post-Lapita sites in accordance with the research prerogatives of the Tongan Dark Age Project initiated at the ANU. He returned to the site at Talasiu (TO-Mu-2) and sampled the midden for ceramic and organic material for radiocarbon dating, reporting on the presence of decorated and plainware pottery within the same context. Most recently, Geoff Clark returned to Tongatapu in 2006–2008 to assess the construction sequences of the monumental sites of Heketa and Lapaha as part of two Australian Research Council (ARC) grants exploring the development of social hierarchy in Tongan prehistory.

These surveys of the archaeological landscape on Tongatapu and subsequent excavations have provided valuable insights into the nature and timing of cultural deposits. The sites were all noted to have intact deposits with little disturbance related to subsequent occupation. The morphology of the material culture and age ranges of dating material (shell and charcoal) also indicated that each of these sites represented a different chronological period within Tongan prehistory spanning over 2000 years. The presence of refuse such as shell, fishbone and charcoal within the deposits at the sites also pointed to the likelihood that the sites were where domestic practices such as cooking and rubbish disposal took place. These three sites were selected for my PhD research based on site preservation and the reasonable possibility that macrobotanical remains might be recovered that would allow archaeobotanical exploration of plant use in Tongan prehistory.

Field methods High-resolution excavation techniques were employed at Talasiu, Leka and Heketa to gain information about subsistence practices and vegetation history at the three sites. Botanical, faunal and artefactual remains were collected from test-pits of around 50x50cm using simple bucket flotation and wet sieving. These techniques are recommended for use in tropical climates (Fairburn 2005b; Pearsall 2010), where access to water and technical equipment can be limited. Some further revision of these techniques was required due to issues that were encountered during fieldwork, and therefore a revised protocol was established that suited the conditions specific to Tongatapu during the 2011 season. The methodology can thus be broken down into four sequential activities:

1. Excavation and bulk sampling 114

2. Bucket flotation 3. Wet sieving 4. Sorting, identification and quantification

Excavation: Test pits of 50x50cm were marked out using string and excavated in 5cm arbitrary levels, referred to as ‘spits’. There was slight variation in the size of the test-pits where contemporary cultural deposits had been already sampled in previous test-pits at particular sites. If a change in stratigraphy was noticed during excavation, a new level was begun. All excavated material was placed into buckets for flotation with a water-proof label. Any observed charcoal concentrations were sampled in situ and wrapped in aluminium foil. The charcoal locations were then recorded on datasheets for each spit. The datasheets also included detailed information upon the soil texture, colour, sorting, and inclusions. Bulk soil samples of around 100g were retained from a 10cm-wide bulk soil sampling column that was measured out and marked on the test pit wall, with samples reserved for microbotanical analysis.

Bucket Flotation: The excavated material from each level was weighed using a spring balance, and the volume was calculated by pouring the sediment into a bucket with volume marks (litres). This measurement was then recorded in a flotation logbook. The material was then divided into buckets, so that each bucket was filled to one-third (33%) of its volume. Water was added until the bucket was nine-tenths (90%) full, and the material was gently stirred by hand. This mixture was left to settle and deflocculate for around 5-10 minutes (after Fairburn 2005b).

A flotation sieve was set up by pegging muslin mesh into a bucket with large holes cut into the base. To recycle water, this rested on two cut branches that sat atop a plastic washing tub. The excavated material was then decanted into the muslin, allowing any material that was floating on the surface of the water to be separated from the heavy residue that had sunk to the bottom of the bucket. More water was then added to the heavy residue, and the process of settling and decanting was repeated at least twice until no more material was visible floating on the surface. The muslin was then unpegged from the lip of the bucket and tied to create a sack that held the ‘flot’ sample with a waterproof label showing the site, test-unit number, level number and date. Finally, these flot samples were placed onto a string line to dry in the sun, and then put into labelled aluminium foil envelopes in zip-lock bags.

Wet sieving: The remaining heavy residue was taken to a new station for wet sieving through a 3mm mesh sieve. Again water recycling techniques were employed by keeping water in two large plastic washing tubs. The sieves were immersed in these tubs to just below the rim and agitated. Keeping the rim out of the water ensured that no larger material floating on the surface (such as leaves) could enter the sieve while being agitated. Once all dirt was removed from the sample, 115 the remaining material was placed onto a plastic rice sack with a water-proof label to dry in the sun.

Sorting, identification and quantification: Some basic sorting of the dried material was carried out in the field. Heavy residue from each level was dry-sieved using a 6mm mesh sieve to create a large (>6mm) and small (<6mm) fraction. Each fraction was sorted and recorded separately. These fractions were placed onto trays and sorted using tweezers into basic material types: ceramics, lithic material, other artefacts, bone, shell, charcoal, seeds and other organics. The sorted fraction were then bagged and labelled. After being returned to the ANH laboratories at the Australian National University, further sorting and quantification of both the heavy residue and the flot samples was carried out. The heavy residue was sorted in much the same way as that in the field, and weights of the various artefacts and material types were recorded. The flot samples were also sorted into material types such as seeds, wood charcoal, charred root and tuber parenchyma, land snails, bone and insect remains. The weights and counts of these were then recorded on datasheets which were then digitised.

Figure 6.1 Map showing location of archaeological sites included in this study from Tongatapu Site descriptions

Talasiu (TO-Mu-2) All three sites were cultural midden deposits of varying sizes and concentrations. Talasiu (TO- Mu-2) in the Mua region is a late Lapita-associated site which was first excavated by Golson 116

(1957). Golson’s field-notes (1957) describe the density of shell and pottery within his excavation units, causing his team to halve the size of the original unit from 10ft² to 10x5ft. During survey, Spenneman (1986:38) noted that the midden site extended as far south as the Langi or monumental stone tombs (J18-J19) but was distinct from another nearby midden (TO- Mu-67). Spenneman sampled the midden and recorded a mix of decorated and undecorated pottery, stone flakes, adzes and adze fragments, and a coral abrader alongside some fragmented human remains.

Evaluation of the topography and geology of the Mua region indicates that the midden was originally deposited on the shore of the Fanga’Uta lagoon; however, later extensive land reclamation using coral rubble, limestone and soil in-filled the inter-tidal lagoon flats, and the site is now approximately 200m inland from the current lagoon shore, except to the north where an inlet cuts close to the site (Dickinson 2007). The midden stretches approximately 100m along the old palaeoshore (Spenneman 1986). The site could have been an attractive location for settlement in prehistory, with a small limestone solution channel and spring located near the site that provides a source of fresh water and feeds into the lagoon (Spenneman 1986; Valentin and Clark 2013). Several cultural strata are present at the site, the earliest of which is composed of many cooking and other domestic features cut or embedded into the reddish sterile clay. Above this is the deposition of a dense midden of near shore and inter-tidal shellfish and fish from the lagoon. Cut into this midden and the earlier domestic features are a number of complete and incomplete burials, believed to be mostly contemporary with the deposition of the midden from 2700-2500 cal BP (Valentin and Clark 2013).

The only test pit excavated at Talasiu (TP2) for this project was a 50x50cm test-unit that sampled the cultural midden, and cut into the sterile basal clay by 5cm. The whole unit was a total of 100cm in depth, with every 5cm of deposit excavated, processed and described as a separate level or spit. All excavated material from these spits was processed using flotation and wet-sieving to isolate botanical, faunal and cultural material. A total of 19 spits were excavated and processed for botanical remains. 100gm bulk samples were taken from each level within a 10cm sampling column marked out on the north-facing profile of TP2, where stratigraphic levels were most distinctive, for starch analysis.

Leka (J17) Langi Leka (also known as Lekamakatuituioha or ‘Puipui’) is a four-tier monumental tomb or langi (J17) built during the classic Tu’i Tonga chiefdom phase of Tongan prehistory, and is said to be associated with Tu’i Tonga Tulunga (McKern 1929:41). The langi is located about 400m inland from the Fanga’Uta lagoon in the community known today as Lapaha, within the Mua region of Tongatapu. Geophysical survey at Langi Leka indicates that the langi had been built on top of an older cultural deposit. Initial excavations were carried out in 2008 to assess the

117 nature and timing of this older occupation through test-pitting. These prior excavations also shed light on the construction sequence for the langi. Attempts were made to relocate TP1which had been abutting Langi Leka. It was believed that TP1 had been found, and TP2 was subsequently excavated parallel to this location further west. A 2x1m test pit was measured out and extended north from the base of the first tier of the langi. The red clay and coral construction rubble debris was recorded and then removed in bulk, so that the darker cultural material below could be excavated and processed in a 100x50cm test pit in the northern end of the excavation. It turned out this unit missed the more dense shell midden that pre-dated Langi Leka by about 5m. Despite this, seven 5cm levels were processed for botanical remains below 90cmbd through flotation and wet-sieving.

In a second attempt, another test 2x1m unit (TP3) was opened up approximately 2m to the east of TP2. TP3 also abutted the langi, and was sampled from 90cmbd within the cultural material using the same techniques employed for TP2, but this time from within a 100x50cm test pit in the southern end of the excavation. Cultural deposits were encountered in both TP2 and TP1, although these were not of the same density noted during previous excavations. TP3 was abandoned and back-filled after excavating to 110cmbd. TP1 was finally located after a system of shovel-test pitting along the base of Langi Leka was employed and the modern fill was quickly removed. The northern baulk was cleaned and chosen for sampling the cultural deposit observed at 95cmbd. A 25x50cm test unit was cut into the baulk, and material above 95cm was discarded. Sampling began below this in the dense cultural midden with every 5cm level excavated, processed and recorded in the same manner as other test pits made at Talasiu and Leka.

Heketa (TO-Nt-2) The final site chosen for excavation and botanical sampling was located in the north-east at Heketa near the ‘esi’ or chiefly backrest known as Makafakinanga (TO-Nt-2) situated close to the Ha’amonga a Maui trilithon (TO-Nt-1). This complex is composed of nine stone structures (Nt-1 to 9) that represent a short period of monumental stone architecture construction, forming an early centre of the classic Tu’i Tonga chiefdom (Clark and Reepmeyer 2014) or state (Kirch 1994). Tongan traditions associate most of the structures with the 11th Tu’i Tonga, Tuitatui, while an earth mound is tied to the 10th Tu’i Tonga, Momo (McKern 1929). Dates for the construction of the stone architecture all fall within the 14th century (Clark and Reepmeyer 2014; Spenneman 2002), but several earlier deposits have been located underneath these structures dating to the 12th-14th century AD, indicating that the site may have also been a significant place associated with early Tu’i Tonga Hikuleo for several centuries before this. In traditions the centre of the chiefdom was moved south to Lapaha by the 12th Tu’i Tonga Talatama.

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This original test pit (TP1) had been excavated in order to assess the construction sequence for the ‘esi or backrest monolith. The aim was to cut into the baulk of this original test pit to sample a cultural midden and charcoal deposit that predates the erection of the ‘esi which had been recorded and dated previously. With this in mind, a 50x50cm test pit (TP2) was marked out about 1.5m away from the ‘esi. The first test pit was excavated to 90cm below datum (bd), when the sterile clay was encountered. Each 5cm level was sampled for botanical, faunal and artefactual material through flotation and wet-sieving. Assessment of the profile and excavated material suggested that this test pit had only sampled the edge of the feature exposed in the original test pit.

Further test pits were then used to locate the looser and more mixed backfill of TP1, and then this backfill was removed to expose the original walls of the test pit. The final unit, TP3, was excavated into the cleaned face of the baulk between TP1 and TP2. TP3 was a 50x30cm test unit that was excavated to 105cmbd below datum, where the sterile clay was again encountered. All material from each 5cm level after 40cmbd was processed using flotation and wet-sieving. The material above this point was all sterile red clay which capped the cultural deposits and topsoil, and had already been sampled within TP2. This strategy allowed specific targeting of the cultural deposit within this test unit.

Stratigraphic descriptions

Talasiu (TO-Mu-2) The whole test unit consisted of very compact shell midden with limited sediment matrix. There was very little if any topsoil, and below this some distinctions can be made within the midden deposit. The top 20cm (Layer 1) below the current surface level contained a disturbed and highly fragmented shell deposit with a light grey brown silty matrix. Charcoal, coral and limestone inclusions were very common. The presence of many small rootlets combined with a lack of protective topsoil contributed to the disturbed nature of this uppermost deposit. Below this was a region of more compact shells to a depth of around 35cmbd (Layer 2) that were also highly fragmented within a coarse yellow brown silty clay matrix. Higher concentrations of charcoal were noted throughout the deposit and possible volcanic ash lenses. The transition between Layer 2 and 3 is not level, indicating that the surface of Layer 3 was undulating.

Layer 3 was a thick (35-60cmbd) deposit of loose large shell fragments with a friable yellow red clay matrix and large amounts of charcoal throughout. More pottery was located between 50-60cmbd, and large fragments of Tridacna were found at the base of this deposit. A small feature (Feature 1) was cut into Layer 3 from above and had more compact shell. It is possible that this was a disturbance from a tree root observed within this deposit. Layer 4 was another undulating layer below this with very compact large shell fragments, charcoal fragments, and a red clay matrix that extends to 80cmbd. At the base of this layer was another 119 possible feature (Feature 2) which was rounded with a diameter of 8cm, and cut into Layer 5. A higher density of shell was noted in the north-west corner of the test unit. The final and bottom layer (Layer 5) of midden was a reddish clay sediment with small crushed shell and charcoal at the base of the midden. The base of the layer slopes towards the north-west corner, and so this was recorded as Feature 3, however this may just reflect the undulating nature of the original ground surface. Importantly, there were also small fragments of dentate-stamped and plainware pottery found inter-mixed throughout the lower half of this assemblage, and near the base of the test unit at 85-90cmbd in the south-west corner. Sterile orange brown clay was encountered at 95cmbd.

Figure 6.2 Stratigraphic diagram of cultural deposits within Talasiu TP2

Leka (J17) All three test pits at Leka revealed an interesting construction sequence for the langi or tomb, with features that were associated with the ditch for slotting the stone facing of the first tier of the tomb into place, and limestone cobbles providing support for these. Below these construction features were cultural deposits associated with occupation at the site prior to the development of monumental architecture at Lapaha. This occupation seemed to be centred near the location of TP4, where the deposit was deepest and contained the highest concentration of archaeological material.

The stratigraphy observed within TP2 represents a range of deposits associated with the construction of the langi, and the older cultural surface upon which the langi was built. The uppermost layer below the surface, Layer 1 was a coral gravel deposit with red brown clay matrix that was more concentrated near the surface and more diffuse towards the base. This 120 layer is most likely construction debris from the langi. Below this was deposit of reddish brown clay (Layer 2) with dispersed charcoal that capped an older cultural deposit (Layer 3). Layer 3 was a medium brown silty clay cultural deposit with many charcoal inclusions. Some red clay from the base of this layer was inter-mixed with the cultural material from 105-125cmbd. This layer sat upon the surface of the sterile red brown clay at 125cmbd.

In TP3 a shallow topsoil deposit (Layer 1) was visible below this surface from 5-15cm below the datum. Layers 2-5 were various coral gravel fill deposits within the ditch cut to allow the placement of the limestone tomb stones. At the base of this ditch was a surface of large limestone cobbles up to 20cm in diameter that were used to hold the beachrock slabs in place. Layer 2 also appeared to extend out to be a coral gravel path that surrounded the base of the langi. Layer 6 was a sterile red brown mixed clay in which was a tephra ash lens (Layer 7). It is believed that the large amount of sediment that created this layer was the result of ongoing tephra ash falls after 1000 BP. These deposits capped the mid-brown silty clay cultural layer (Layer 8) with small dispersed shell fragments and charcoal. This deposit was the only layer sampled for flotation and microbotanical analysis.

The stratigraphy of TP4 resembled that seen within TP2 and TP3, although the cultural deposit below 95cmbd contained a more dense concentration of shell, bone and charcoal. Similar to TP2, this cultural deposit was located below various mid-reddish brown clay deposits (Layers 1-3). The top of the cultural deposit was a mid-brown silty clay layer (Layer 4) from 95cmbd, with some dispersed shell, charcoal, fishbone, fire-cracked rocks and small limestone cobbles. Shell midden composed the lower half of the cultural deposit (Layers 5-6) from 125cmbd, and had internal stratigraphic differences. A mid-brown silty matrix was observed at the top (Layer 5), above a more red clay matrix at the base (Layer 6). The frequency of charcoal within the deposit increased between 135-140cmbd. By 140-145cmbd the cultural deposit began gradually transitioning into red clay with fewer shells and charcoal. The base of the excavation was sterile orange brown clay (Layer 7).

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Figure 6.3 Stratigraphic diagram of cultural deposits within Leka TP2

Figure 6.4 Stratigraphic diagram of cultural deposits at Leka TP4

Heketa (TO-Nt-2) The initial test pit at TO-Nt-2 (TP2) revealed the edge of two deposits, one possibly related to the construction of the ‘Esi, and the other deriving from a pre-stonework phase of occupation at the site. Below the topsoil within this unit was a layer of red brown volcanic clay to a depth of 122 around 20-25cm below datum (bd) (Layer 1). Below this was a layer of beach-rock sand debris inter-mixed with some of this red brown clay to 25-30cmbd (Layer 2). A dark brown clay deposit with some humic material was beneath this to 30-35cmbd (Layer 3), and capped the cultural material. Layer 4 was a deposit of cultural material with dark brown clay matrix and charcoal fragments which was observed below these surfaces. The density of cultural material in this deposit was sparse. The base of this feature was sloping towards the south, and so the deepest point of the cut was at 70cmbd in the western-facing profile at the south-east corner. The shallowest point of the base of the deposit was at 53cmbd. A small pocket of shells followed the contour of the base of this deposit, about 3cm above the sterile clay. This deposit was possibly cut into another older occupation layer that had higher concentrations of shell, coral and charcoal and thus appeared darker from 70-93cmbd within the profile (Layer 5). The base of this deposit grades to less shell and more charcoal at the base. Both cultural deposits sat on top of a surface of sterile orange brown clay (Layer 6). This test pit was dug to approximately 99cmbd when the sterile clay was encountered.

The profile of stratigraphy within TP3 mostly resembled that of TP2. Red brown volcanic clay below the topsoil extended to a depth of 25cmbd (Layer 1). Below this was the same beach-rock sand inter-mixed with this clay (Layer 2), most likely related to the shaping of the limestone slab for the ‘esi. This layer of debris capped a cultural deposit (Layer 3) at 30- 35cmbd. The cultural deposit consisted of dense large shell and charcoal fragments with a dark silty loose clay matrix. There was some tree root disturbance within the deposit in the southwest corner, from 45-60cmbd. Near the base at 85cmbd was a white ashy lens (Layer 5) within a deposit of concentrated burnt crushed shell, charcoal, large whole shells, and many diodont or puffer fish spines (Layer 4) from 80cmbd to the base of the feature. Very high densities of mussel shell and a whole Lambis was located directly underneath Layer 5, grading to higher concentrations of Diodontidae spines in with medium orange brown clay from the base of the layer. The base of the deposit was encountered at 95cmbd, where the feature was cut into the orange brown sterile clay seen in TP2 (Layer 6).

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Figure 6.5 Stratigraphic diagram of cultural deposits within Heketa TP3 AMS dating of cultural contexts In order to identify samples for dating, the wet-sieved, flotation, and in-situ charcoal samples were sorted into those that could be identified as coconut, other endocarp or nuts, and other charcoal (wood and parenchyma). Coconut endocarp was identified based on the surface texture, which consists of a ‘cross-hatched’ type of pattern where clusters of three or four elongated parenchymatous cells are orientated at 45 degree angles to each other, as well as the thickness and density of the cross-section. Coconut is also very difficult to fracture, due to the density of the endocarp, but when it does fracture it does so relatively evenly and cleanly. The methods employed to extract macrobotanical remains from these sites were successful in producing small amounts of coconut from the wet-sieved material and in situ samples at all three sites. In order to develop a chronology for these sites, three samples of charred coconut endocarp were selected from spits (levels) located near the top, middle and bottom of one test- pit from each site for dating at the Waikato Radiocarbon Dating Laboratory (WRDL).

The results of AMS dating indicate that all three sites are from separate and relatively discrete time periods within Tongan prehistory. The midden at Talasiu (TO-Mu-2) can be dated to around 2750-2650 cal BP, and appears to have accumulated over a period of around 100 or less years. All three radiocarbon samples had narrow age ranges that overlapped very closely with one another, indicating a relatively short depositional period. This site therefore represents refuse from a settlement that is on the cusp of late-Lapita and Ancestral Polynesian Society (APS). These dates are supported by the ceramic assemblage from Talasiu, which is composed primarily of plain ware with a few dentate-stamped sherds in lower levels of the midden (Clark et al. In press; Golson 1957; Spenneman 1986).

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Figure 6.6 Calibration of radiocarbon dates from Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2)

Occupation underneath Langi Leka (J17) can be dated to around 1300-1000 cal BP, and so falls within the Formative Period (1550-750 BP) in Tongan prehistory (see Chapter 1), of which little is known (Burley 1998:365-8; Davidson 1979). The abandonment of pottery traditions around this time leaves an impoverished archaeological record, with a total of only 16 sites on Tongatapu representing these 800 years of occupation. The cultural deposit sampled for radiocarbon dating represents another relatively short period of deposition, with all three date ranges overlapping within a 100-200 year period.

Finally, the sampled Heketa (TO-Nt-2) material was from two deposits. One is around 800-600 cal BP and at the end of the Formative Period, while the other is a later deposit that is related to the development of early monumental architecture in Tonga during the Classic Tu’i Tonga chiefdom after 600BP (Burley 1998:368-79; Clark and Reepmeyer 2014). Two radiocarbon samples were from this lower and earlier deposit, and the remaining sample originates from the more recent feature and fill that caps this older pre-stonework architecture surface.

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Chapter 7 Laboratory Methods

A combination of micro- and macrobotanical techniques was implemented to research and identify plant use at three stages in Tongan prehistory. Sites were selected based on the results of previous archaeological investigations on Tongatapu and sampled for botanical remains using current archaeobotanical methods, modified for tropical environments. A combination of flotation, wet-sieving and bulk stratigraphic sampling was used to extract all micro- and macrobotanical material from each test-pit. Laboratory processing of these botanical remains built on current techniques after experimentation revealed issues with contamination in the laboratory environment and also methods for the dispersal of microcharcoal concentrations within archaeological sediments, and the isolation of starch during heavy liquid separation. A revised protocol for starch extraction is published here for future use in microbotanical analysis. The extracted starch and parenchyma was analysed using both light microscopy and SEM, but was then classified to taxa using a combination of multivariate statistical analysis in the form of Discriminant Function Analysis, identification flowchart keys and visual checking of matches that is outlined in Chapter 8

Microbotanical analysis: Starch residues

Experimentation with starch extraction techniques As emphasised within Chapter 3, a number of previous technical studies of starch residue extraction and processing from sediments have highlighted issues with current protocols, such as sampling strategies, contamination and the use of destructive chemicals (Barton et al. 1998; Coil 2003; Crowther 2009; Korstanje 2003; Parr 2002; Therin and Lentfer, in Torrence 2006b; Torrence 2006b; Torrence and Therin, in Torrence 2006b). New protocols are constantly developed to address the effects of different taphonomic, environmental and laboratory conditions. In the current study, several experiments were designed to solve issues that arose during the first round of laboratory processing, especially potential contamination from the chemicals and equipment used. This experimentation sought to deal with this potential contamination in the laboratory setting, as well as issues with the removal of micro-charcoal that can inhibit viewing of starch granules on glass slides and the recycling of heavy liquid. The results of these experiments enabled the development of a revised protocol that suited the nature of the archaeological deposits being sampled and the laboratory equipment available at the ANU.

Experiment One: Potential for starch contamination from Calgon, de-ionised water, filter mesh, LST and Glycerol The risk of contamination during post-excavation processing has been outlined in previous research by Loy and Barton (in Torrence 2006b), Laurence (2013) and most recently Crowther (2014). Experiments were designed to investigate the potential for contamination from materials

126 used to process the archaeological sediments. To do this, each material was independently tested for the presence of starch, and any observed starch was recorded and counted.

Calgon Firstly, powdered Calgon or sodium hexametaphosphate was tested by mixing the powder with de-ionised water to produce four 500ml samples of 5% Calgon. A magnetic mixer was added to each of the samples, and these were then placed on a hotplate set to 20˚C to allow the Calgon to dissolve into the de-ionised water. Once this had occurred, the samples were each filtered through a 10µm laboratory-grade-filtration mesh. The mesh was then cut into three parts, mounted onto slides, and covered with a cover-slip. Mountant was not added to the samples, as these were constructed only as temporary slide mounts. The slides were then viewed using light microscopy in both brightfield and polarised light.

The results of this experiment suggested relatively high numbers of starch contaminants from powdered Calgon. Sample 1 had a total of 35 grains, Sample 2 contained 12 grains, Sample 3 had 69 grains, and Sample 4 had 37 grains. The predominant starch types encountered were taxonomically identified as wheat (Triticum spp.), and maize (Zea mays). It is possible that contamination of the Calgon occurred in the laboratory storage for the Department of Archaeology and Natural History, as the bag had been left open for a time. It is also plausible that some contamination can occur even within laboratory-grade Calgon during manufacture, as many companies produce both Calgon and powdered maize (along with other dehydrated or milled plant products).

To resolve this issue and enable Calgon to be used as a deflocculant during processing of the archaeological samples, the 5% Calgon/de-ionised water mix was filtered through a 5µm laboratory filtration mesh to remove any potential starch contaminants. The filtered mix was then always covered unless being added to soil samples in starch extraction processes to break down clay particles.

De-ionised water Four 500ml samples of de-ionised water were processed in a similar manner to the Calgon. The beakers were poured through a circle of 10µm filtration mesh and the filter paper was then cut into three parts and mounted on slides as temporary mounts. No starch was observed within these samples during light microscopy, and so it was concluded that de-ionised water could not be a source of starch contamination when used in extracting starch from archaeological sediments.

Filtration mesh Four circles of 10µm mesh laboratory-grade filtration mesh were dampened with de-ionised water, after previous experimentation revealed that this was not a potential source of starch contamination. These circles were then each cut into three parts, placed onto slides, and then

127 covered with a coverslip. Light microscopy revealed that very little starch can be found on filtration paper, with only one to two grains observed on any of the samples (Sample 2), and therefore these are also not a significant source of contamination during processing of archaeological sediments.

Lithium hexametaphosphate (LST) Two samples of unused and two samples of recycled LST heavy liquid at 2.0sg were also tested for starch contamination. These were filtered through individual rounds of 10µm filtration mesh, and cut into strips before being placed on slides as temporary mounts. These slides were then observed using light microscopy and any observed starch morphotypes were recorded. No starch was observed within any of these samples, indicating that these are not sources of starch contamination during processing.

Glycerol Glycerol was the mountant chosen to create permanently mounted slides for both the comparative collection and the starch extracted from archaeological sediments. It has a high refractive index and ensures starch is preserved and protected from enzymatic attack, while also allowing starch granules to be rolled. To test whether the Glycerol used in the Palynological Laboratory in the Department of Archaeology and Natural History could be a potential source of starch contamination, four slides were produced with 100µl of Glycerol on each. These slides were covered with a cover-slip, and then observed using light microscopy. Three wheat starch granules were found in total on these four slides, and so this material is a potential source of starch contamination, but the contamination is very small.

Conclusions These experiments assessed the amount and types of starch within a number of standard or regularly employed chemicals and materials used to extract starch residues. Of the five materials tested, only three contained any visible starch. Glycerol and filtration paper had very small amounts of starch, with less than three granules observed in any one sample. Based on these results it was decided that the use of glycerol and filtration paper within the current protocol was acceptable and required no further processing. In contrast, the test involving unfiltered Calgon highlighted that this material could be a significant source of contamination from modern wheat and maize starch. Up to 69 starch grains were observed in each of the four samples. These findings indicate that a major revision of the deflocculation process was required to enable archaeological sediments to be processed without the risk of contamination. In light of this, it was decided that the pre-mixed Calgon needed to be filtered through a 5µm filtration mesh prior to use. All other materials were acceptable for use within the revised laboratory protocol.

Experiment Two: Removal of charcoal from samples The soil samples collected from Talasiu, Leka and Heketa were taken from shell middens of various densities, and it was noted that there were high concentrations of microcharcoal present

128 in the soil matrix. Microcharcoal can make light microscopy difficult by obscuring the visibility of any potential starch grains, and so needs to be either removed or dispersed in starch extraction. Several experiments were carried out to establish whether particular methods were effective in this, and also to gauge the effect these methods had on starch residues. These replicated and built on experiments carried out by Crowther (2009:62-82).

Lithium hexametaphosphate (LST) One method tested for the removal of charcoal was to use Heavy Liquid to remove light fractions of material from soil samples. Wood charcoal has a specific gravity (sg) or ratio of density of 0.4sg, which enables it to float on water during archaeobotanical flotation techniques. However, the microcharcoal observed in this experiment has instead settled in water during Step Two of the methodology outlined here. It was hypothesised that this microcharcoal may be able to be removed using a heavy liquid with a specific gravity heavier than water (1.0sg), but lighter than the specific gravity of starch (1.7sg).

One archaeological sediment sample (Talasiu Spit 18) was selected to be sub-sampled and processed to the point where Heavy Liquid in the form of lithium heteropolytungstate (LST) was added using the methodology outlined below. Thirty millilitres of LST at 2.0sg was added to the sample, and centrifuged at 1500rpm for 30mins. This was decanted through a 5µm filtration mesh, enabling the LST to be separated from the extracted material and thus easily recycled. The mesh was then placed in the top of a Falcon tube and de-ionised water was used to wash the material caught on the mesh into the tube. All material lighter than 2.0sg and larger than 5µm was therefore retained. Another LST solution at 1.2sg was then added, and the process repeated so that a fraction that was lighter than 1.2sg was separated. The heavy fraction was also retained as this should contain any starch residues.

The results of this experiment indicate that microcharcoal cannot be separated using LST set at 1.2sg. Most of the charcoal examined tended to become waterlogged when exposed to liquid over a sustained period of time, and became too dense to float. The differential specific gravities of microcharcoals are probably a result of differing wood densities before charring, particle size, and density as a result of water absorption. These changes probably occur either in the soil or during Step One and Step Two.

Hydrogen peroxide (H2O2) The second method tested in an attempt to remove charcoal from the archaeological sediments was the addition of hydrogen peroxide or H2O2. Talasiu Spit 18 was again selected to be sub- sampled for this experiment. The sample was processed until the point where it was reduced to allow slide preparation. The sample was placed into a thin glass centrifuge tube under the fume hood, and 10% hydrogen peroxide was added to the sample one drop at a time to gauge any chemical reaction. There was no immediate reaction, nor to a higher percentage dilution of 30%

129 added in the same manner. There was only a very small reduction in the amount of charcoal observed within the sample.

The final test in this experiment involved the application of 30% hydrogen peroxide and then the sample was placed in a beaker of water on a hotplate to stimulate a stronger chemical reaction. The water was warmed to 30˚C, and the sample was observed for 1.5 hours within which time there was still no reaction. A slide was prepared after the final test in this experiment to observe any effects of these chemicals on starch morphology. It was clear that most starch had either gelatinised or dissolved. Very low numbers of starch in native condition were observed in this test sample.

Simple dispersal The final method tested in this experiment was not to remove the charcoal, but to allow simple dispersal within the sample. Rather than preparing only one or two slides from the reduced samples after processing, three to four slides were constructed. This allowed any micro-charcoal present in the samples to be dispersed over a larger number of slides, reducing the possibility of these charcoal particles obscuring any starch granules present in the samples.

Conclusions The use of chemical agents to remove microcharcoal from archaeological sediments was not successful for a variety of reasons. Heavy Liquid separation using LST at 1.2sg was not able to float off micro-charcoal, due to the varying densities of wood charcoal. Some microcharcoal was removed during this process, but high concentrations remained in the heavy fraction that also contained the extracted starch grains. Similarly, 10% and 30% hydrogen peroxide proved to be ineffective in dissolving charcoal, even when heat was added in an attempt to produce a stronger reaction. In addition, such concentrations of hydrogen peroxide affected starch morphology and preservation, rendering the use of these chemicals unacceptable.

Experiment Three: Removal of supernatant and recycling of heavy liquid A simple experiment was carried out during the first phase of laboratory processing of the archaeological soil samples to allow the separation of the supernatant after heavy liquid in the form of lithium heteropolytungstate (LST) was added. Current methodologies (Horrocks 2004; Therin and Lentfer, in Torrence 2006b) either pipette or decant the supernatant into a separate tube, leaving the heavy residue behind in the original centrifuge tube to either be kept for a second phase of heavy liquid separation or to be discarded. Experimentation was conducted to ease recycling of heavy liquid for use within other samples and reduce inter-sample contamination.

During this experiment six samples from one spit (level) at Talasiu (TO-Mu-2) were processed up to the stage prior to heavy liquid separation. Exactly 30ml of LST at 2.0sg was added to all four of these samples within centrifuge tubes which were then placed into the

130 centrifuge and spun at 1500rpm for 15mins. At this point two samples were chosen to attempt pipetting off the supernatant, two samples were selected for decanting, and two samples were chosen to test a new method which involved decanting the supernatant through a 5µm mesh (Janelle Stevenson, pers.com). The reason for attempting this new method was to reduce the amount of non-starch material within the sample after heavy liquid processing and to ease the process of recycling the heavy liquid through separating the starch from the LST immediately. Each set of samples were then processed accordingly. Individual techniques for separating the supernatant were assessed for ease of use in terms of the time it took to carry out the separation, the efficiency of the separation in terms of the amount of heavy residue within the supernatant, and the subsequent steps needed to remove and recycle the heavy liquid.

Pipette Technique Pipetting the supernatant off the top of the heavy residue which had sunk to the bottom of the centrifuge tube proved to be a very efficient technique for separating the light and heavy fractions. Very little of the heavy residue was accidentally added to the light fraction. However, the method was also very time-consuming as only small amounts of the supernatant could be removed at a time, taking around 5 minutes. The heavy liquid then needed to be removed from the light fraction through various steps involving the dilution of the sample to wash the heavy liquid and retain starch residues. The heavy liquid then needed to be passed through a glass mesh to remove any contaminants. To increase the specific gravity of the heavy liquid, it was put inside a glass beaker on a magnetic stirrer at low heat and monitored until enough water had evaporated to return the LST to 2.0sg. This was again a time-consuming process, taking around eight hours to enable the heavy liquid to be reused.

Decanting Technique Decanting the supernatant into a new centrifuge tube took less time to separate the heavy and light fractions—approximately 1 minute— but was less effective. There was some mixing of the two fractions as the last of the supernatant was carefully poured into the second tube. This meant that the final slides had more organic material than those made from the pipette samples. The same process was then used to recycle the heavy liquid through dilution, sieving and evaporating water to return the LST to 2.0sg.

Sieving Technique Pouring the supernatant through the 5µm mesh was a time-consuming process as small amounts of organic residue from the top of the heavy fraction was inter-mixed with the light fraction, as seen within the decanting technique, and combined with charcoal to clog the mesh. A pump had to be used to slowly allow the supernatant to pass through the mesh, taking around 10 minutes. The light fraction remained on the 5µm mesh, and was then put into the top of a centrifuge tube and washed off into the tube using de-ionised water. Only one wash was required to remove any residual LST from the sample before the sample could be reduced for slide construction.

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Despite being more time-consuming in terms of separating the supernatant from the heavy fraction, the technique was very effective at separating the heavy liquid from the light fraction quickly without any dilution, enabling the LST to be used immediately after only one further step of sieving through a glass filter paper.

Conclusions Both pipetting and decanting the supernatant off from above the heavy fraction were more time- effective methods than the sieving technique in terms of separating the light fraction for further processing. Of these, pipetting was the most effective technique to reduce the amount of heavy residue accidentally entering the light fraction during separation. Once these samples were separated, both the pipette and decanted samples required further processing to remove the heavy liquid and enable it to be re-used for subsequent samples. This processing added approximately eight hours (dependent on the amount of heavy liquid being recycled) to the laboratory protocol. In contrast, the sieved samples only involved one step of centrifuging and decanting that took around four minutes, before the sample was ready to be reduced and placed on a slide. The process of removing contaminants from the heavy liquid by filtering through a glass mesh, and repeating this step added another five minutes to the protocol before the heavy liquid was ready to be reused. Therefore, despite the initial processing time, the sieving technique is the most time effective and efficient overall method for separating the supernatant and recycling the heavy liquid.

Laboratory processing: Revised starch extraction protocol After experimentation highlighted potential sources of starch contamination within the materials used during standard starch extraction processes, a method for dispersing charcoal and a new technique for removing the supernatant after density separation and recycling heavy liquid were incorporated into a final extraction protocol. This protocol was based on modification of techniques published by Horrocks (2004), Torrence (2006a, b) and Field (pers. comm. 2012). It involved steps to sub-sample, deflocculate, settling, sieving, centrifuging, density separation, and reduction of samples for slide construction.

Note on sub-sampling and site variation Talasiu was the first site to be analysed for the presence of starch residues. A 3gm sub-sample was taken from each 5cm spit excavated from the test-pit, and processed to extract preserved starch residues. After radiocarbon dating revealed that all three sites represent relatively discrete time periods, it became clear that there was little intra-site age variation and so few chronological differences would be observed within each of these sites. A decision was made, therefore, to sub-sample every second 5cm spit to gain a representative distribution of starch preservation within the deposits sampled at Leka and Heketa.

Heketa had very low quantities of starch during the first round of analysis. As a result, a decision was made to re-process four spits from the site with new sub-samples. A few changes 132 were made in the processing technique to assess the potential of different variables in methods for starch extraction. The samples were sieved through a 250µm filtration mesh, rather than 125µm. Secondly, the samples were placed in the centrifuge for 30 minutes rather than 15 minutes during heavy liquid separation. This gave a larger window of time for starch to release from the heavy fraction and become part of the supernatant.

Laboratory protocol for starch extraction from archaeological sediments  Step 1: A 3gm representative sub-sample of each archaeological sample was measured out and placed into a 50ml capacity Falcon centrifuge tube. Ten millilitres of filtered 5% Calgon mix was then added as a deflocculant to each of the tubes before the tubes were capped and individually vortexed for 30 seconds.  Step 2: The samples in the tubes were poured into a glass beaker with 500ml of de- ionised water and covered. Stokes Law was employed to calculate the amount of time that the samples should be left to allow organic particles to settle. It was established that samples should be left for at least three hours to allow the clay particles to separate, and organic spherical particles larger than 5µm to settle 10cm at terminal velocity in water at temperatures of 17-21˚C. After three hours each sample was decanted, refilled with 500ml de-ionised water and covered. These processes were repeated at least twice or until the supernatant was clear.  Step 3: The sample was then poured through a 125µm sieve to remove the larger organic fraction.  Step 4: The remaining sample was poured into a Falcon tube and capped, before being centrifuged at 2000rpm for four minutes and then decanted. This process was repeated until the supernatant was visibly clear.  Step 5: As much liquid was removed at this point and then 30ml of heavy liquid in the form of lithium heteropolytungstate (LST) at 2.0sg was added to each sample. The samples were then again put back into the centrifuge and spun at 1500rpm for 15 minutes.  Step 6: Each sample was decanted through a new 5µm mesh circle, enabling the LST to be separated from the extracted material and thus easily recycled. The mesh was then placed in the top of a Falcon tube and de-ionised water was used to wash the material caught on the mesh into the tube. All material larger than 5µm was therefore retained.  Step 7: The samples were then centrifuged at 2000rpm for four minutes and decanted. This process was repeated twice to wash and dispose of any remaining LST.  Step 8: The final step involved reducing the samples to less than 5ml by allowing the samples to settle overnight and then pipetting off the supernatant. This remaining material was placed onto slides and allowed to dry before Glycerol was added. The material on the slide was gently agitated to release the dried starch residues from the

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glass, and then a cover-slip was placed over it and sealed with nail polish to ensure that the slide was protected from contamination.

Light Microscopy All samples of extracted starch residues from Talasiu, Heketa and Leka were observed and imaged using light microscopy. A Leica DM6000 Compound Transmitted Light Microscope within the Centre for Advanced Microscopy at the ANU was used to view starch in both brightfield and cross-polarised light. Each slide was viewed in horizontal transects, enabling all starch to be observed and eliminating the prospect of accidentally recording the same grain twice. Images were taken of every starch grain encountered, with quantification capped at 100 grains per sample. These 100 grains were usually counted over several slides from a particular sample, meaning no sampling technique was employed. Often the samples contained less than 100 grains in which case absolute counts were recorded. Images were taken in both brightfield and cross-polarised light to enable recording of attributes related to the size and shape of the grain, and also the extinction cross which can only be viewed in cross-polarised light. Each imaged grain was given a number which correlated with the file names for any digital photographs taken of that grain. The grains were rolled by tapping the top of the cover-slip gently to enable accurate description of the three dimensional shape. This technique also allowed the observer to note if the grain was lenticular or biconvex in one plane of view.

Archaeological starch classification: Assemblage-typology approach To explore morphological patterning in the archaeological starch assemblage, a number of starch morphotypes were created based on a combination of the nominal and metric attributes. This analytical technique has been used elsewhere as a means of gauging morphological variation within archaeological assemblages (Barton 2005; Crowther 2009:150-2; Lentfer et al. 2002; Mercader et al. 2008). The technique is most often used where a comprehensive comparative collection is not available and therefore the ‘rules’ of group membership are not clear (Crowther 2009:150). Breaking down the archaeological assemblage into ‘types’ enabled the range of morphological diversity within each context to be studied, and provide an estimate of the number of taxa present. This approach was also useful to identify starch that had been modified through various cultural and natural taphonomic processes, using the work of Barton (2009), del. P. Babot (2003:69-82) and Crowther (2009:19-61, 2012). The primary morphological variable used in this study was three-dimensional shape, followed by shape modifiers such as the number and type of pressure faceting, and the presence and type of fissuring at the hilum.

Pinpointing the species from which these starch types could have originated was a critical next step. To do this the morphotypes were compared to those observed within the comparative collection. This was done by analysis of the morphological patterning recorded

134 within the Excel database, and images taken during light microscopy and SEM. Most often, a number of matches were possible. To narrow this list down further, the archaeological starch types were then compared to the length and width ranges of the matches. This assemblage-based approach can also highlight the possibility that archaeological starch does not originate from any taxa in the comparative collection, and therefore may only be identified to genus or family level, or remain as an unknown morphotype.

Archaeological starch classification: Multivariate statistical analysis After the assemblage typologies had been created and some basic taxonomic identifications were given to the extracted archaeological starch based on visual or size characteristics, multivariate statistical analysis was carried out to further extrapolate or confirm these identifications. The extracted archaeological starch was classified using multivariate statistical analysis in the form of Linear Discriminant Function Analysis (DFA). The statistical analysis was based on the analysis of the reference material, and was used to classify archaeological starch using the same variables. These were classified in the same manner as the reference starch, but as ‘ungrouped specimens’ that were not included in the learning parameters for the DFA.

The database for each test-pit from each site was divided into two based on the orientation ratios, and the range of variables altered according to those assessed and deemed to be significant from the analysis of the comparative collection. Instead of being given a ‘grouping’ as within the datasets from the comparative collection, a ‘?’ was entered into this column for each archaeological starch grain. Each archaeological starch grain was named in the ‘point’ column according to the spit from which they were extracted, and the grain number given to them during recording. In this way, each archaeological dataset could be individually compared to one of the two datasets (side-on or end-on) from the comparative collection, and each starch grain given a predicted classification alongside the reference starch. These were given through the ‘classifier’ output for each dataset, and each starch grain could be traced back to the sample from which it was extracted through the name it was given in the point column.

Quantification of the classifications was carried out so that the number of starch grains matched to each species within a particular sample could be calculated. These figures gave an overview of the range of species potentially present within each sample and the quantities of starch classified within these groupings. During this phase, the results of classifications from both datasets were combined; however, it became clear that in order to do this some system was needed to filter the classifications, creating a statistical gauge for the level of confidence in the presence of any species within a sample.

Confidence in these classifications in a particular sample was made by fulfilling a number of criteria. High confidence classifications had to have a successful reclassification rate 135 within DFA for that species of over 60%, and more than five grains had been matched to that species within that sample. Moderate confidence classifications had to either have over 60% correct reclassification but less than five grains matched, or less than 60% correct reclassification and over five grains matched to that species within that sample. Low confidence classifications were given when the reclassification for that species was less than 60%, and less than five grains were matched to that species. In this way, each classification made using DFA was acknowledged, but a means of gauging the actual likelihood of that species being present in a particular spit was also provided. The identifications were then confirmed using visual checking of the images of the archaeological starch. These were compared with the images of the species taken using light microscopy and SEM to which the DFA had classified each grain, confirming or eliminating these as possible taxonomic classifications.

Macrobotanical analysis: Charred parenchyma and endocarp

Laboratory analysis Macrobotanical remains were collected using three different methods during the excavation process. Any large charcoal fragments above 1cm in diameter observed during excavation of the test units were collected and placed in aluminium foil envelopes. These samples were labelled according to the site, test unit and spit from which they derived. Charcoal and seeds were also collected within both the light and heavy fractions during flotation. The light fraction floated during bucket flotation when water was added to the soil, and the heavy fraction remained in the bottom of the bucket to be wet-sieved. Some charcoal can become water-logged and therefore sinks during this process. This depends on the degree of charring within the fragments, and porosity of the material.

In the quarantine laboratory at the ANU, the heavy fraction was sieved through a 3mm mesh to create two size fractions, smaller and larger than 3mm. These fractions were then sorted into material and artefact types including shell, ceramics, land snails, seeds, and charcoal. These were bagged separately and labelled according to the site, test pit, spit and sample type (in situ, flot or wet-sieved). The flot (light fraction) was likewise sorted into material types such as charcoal, seeds, insect remains, pumice, small bone and landsnails. The charcoal from each of these sources, including in-situ, was then further sorted into parenchyma, wood charcoal and endocarp.

Charred endocarp Where possible the endocarp was identified to genus or species level. This taxonomic identification was carried out first through the observation of surface morphology and features, and second through fracturing endocarp fragments down the radial section, and viewing the cell arrangement. Coconut (Cocos nucifera) has a distinctive surface texture on both the exterior and interior surfaces of the endocarp. This texture is created by a cell arrangement that creates a

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‘cross-hatched’ effect through long, thin parenchyma cells that stack horizontally and vertically alternately. Candlenut (Aleurites moluccana) is identifiable through distinctive surface morphology. The exterior surface of the endocarp has a bumpy and slightly warped texture similar to a walnut, and a high degree of curvature relative to fragment size.

Parenchyma Parenchyma was identified to taxonomic level using the range of morphological features recorded within the comparative collection. Each fragment tentatively identified as parenchymatous was firstly described in terms of general and surface morphology (Hather 2000), and then fractured along the transverse and longitudinal planes to record cell arrangement and bundling features. The fractured parenchyma was placed into a small Petri dish and nestled in a bed of salt, and an Olympus Compound light microscope was used to view and image the fragments in reflected light. These fragments were confirmed as parenchyma based on morphology which differs from wood charcoal. These morphological traits include a more rounded surface due to the exposure and erosion of cells; surface features such as buds, detachment scars and spines; few visible rays and thus a more uneven fracture; consistent cell shapes that are usually rounded or angular; and the presence of distinctive vascular bundles or tissues.

Each fragment was identified as either stem or root-derived vegetative parenchyma, and then to species of origin where possible. The identification flowchart key was utilised to provide either a single identification or a list of possible identifications where no further breakdown of the taxa is possible due to morphological overlap, or other features of the archaeological parenchyma are not visible. These taxonomic identifications were then confirmed using images from both light microscopy and SEM. The images could also be utilised to eliminate or confirm taxa where a list of possible identifications is provided.

Classifications were then given based on the determination system outlined in Chapter 5 using identification criteria and associated levels of confidence compiled by Paz (2001) and by Oliveira (2008). This determination system has been used successfully by these researchers in the past to analyse macrobotanical plant remains from a range of archaeological contexts in the Asia-Pacific region (Barker et al. 2011; Barton and Paz 2007; Oliveira 2008; Paz 2001, 2005). The identification key is useful as a means of breaking down the morphology in the comparative collection. However, there was a reasonable likelihood that some samples would not match any of the specimens within the comparative collection due to both the condition of fragments and also the fact that the reference collection is not exhaustive. The reference collection also has a general focus on economic and non-economic plant taxa from Western Polynesia, and so would require the inclusion of additional specimens for use in other geographic locations.

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Chapter 8 Results

Presentation of results is divided into four main sections. The first section considers the quantification and identification of macrobotanical remains from all test units and sites. The various extraction techniques utilised in the field is also compared in terms of the effectiveness of the recovery of charred endocarp, wood charcoal and parenchyma. Each site will also be compared in terms of the quantities of identified material, the distribution of these in test units and the results of a case study presented for the identification of parenchyma at Talasiu. The second section focuses on the quantification and identification of microbotanical remains in the form of starch grains. Once again each site is compared in terms of starch preservation, the numbers of identified species and the distribution of starch within test units according to dated cultural deposits. The third section analyses a number of Western Pacific plant production systems using theoretical insights from Evolutionary and Human Ecology for assessing efficiency of yield and associated nutritional returns. These modern examples provide a useful range of models within which the past systems, represented by archaeobotanical remains, can be assessed and placed in the final section of this chapter.

Macrobotanical analysis

Quantification of charcoal Macrobotanical assemblage breakdown Overall, macrobotanical preservation within all of the sites and test units was relatively good, with small quantities of a range of plant material recovered. However, these remains only composed up to 0.46% of the total pre-processing material weights removed from these units (including soil, shell and other matrices). Within these assemblages, the largest quantities of macrobotanical material were recovered from Talasiu TP2, with a total weight of 60.8g. The smallest assemblage was recovered from Leka TP3, however only three spits were processed from this test unit.

The quantification of the macrobotanical assemblage was primarily based on the separation and analysis of three main components: coconut, other endocarp, and wood charcoal, from which vegetative storage parenchyma was separated later. Wood charcoal composed the bulk of the macrobotanical assemblages from almost all the test units at Talasiu (TO-Mu-2), Leka (J17) and the Heketa (TO-Nt-2), making up around 78% of the collective weight (see Table 8.1 and Figure 8.1). Only Leka TP4 varied from this trend, with wood charcoal making up only 26% of the macrobotanical assemblage from this test unit. In particular, Talasiu TP2 had the greatest amount and distribution of non-endocarp charcoal (53.4g). For this reason, this test unit was selected as a case study for the separation and identification of vegetative storage parenchyma. Only very small quantities of parenchyma (<1g) were found in any given sampled deposit. Aside from wood charcoal, non-coconut endocarp was the second most common 138 macrobotanical material recovered and quantified, composing around 17% of the total assemblage. Coconut endocarp was recovered from every test unit, but only in small quantities and so made up only 5.8% of the overall assemblage.

The quantification of macrobotanical materials is analysed here in terms of the rates of recovery from flotation, wet-sieving and in situ collection, and the overall quantities, as well as the distribution of these quantities within each test unit. This quantification indicated both the success of various methods for extraction of particular plant remains, as well as the effects of site taphonomy and formation processes upon archaeobotanical assemblage composition.

Botanical material Talasiu Leka TP2 Leka TP3 Leka TP4 Heketa TP2 Heketa TP3 Total (g) % of assemblage Coconut (g) 1.6 0.6 0.1 0.7 1.6 1.2 5.8 4.832125302 Endocarp (g) 5.8 0.3 2.3 4.1 2.6 5.1 20.2 16.82912605 Wood charcoal/parenchyma (g) 53.37 8.47 2.4 1.72 4 24.07 94.03 78.33874865 Total (g) 60.77 9.37 4.8 6.52 8.2 30.37 120.03 % of total material weight 0.0219 0.0064 0.0065 0.0696 0.1708 0.4658

Table 8.1 Summary of total macrobotanical assemblages from all sites and test-pits

Figure 8.1 Composition of overall macrobotanical assemblage in terms of abundance

Coconut endocarp Basic sorting processes highlighted an interesting pattern in the macrobotanical extraction process. Coconut endocarp was extracted exclusively during the wet-sieving stage of archaeobotanical processing in the field. This is most likely because of the higher density of coconut endocarp, which does not float as well as wood charcoal, or other varieties of endocarp. Another possibility is that coconut endocarp does not disaggregate easily from clay soils during the initial deflocculation process before flotation. No experimental tests were carried out to test either of these ideas. These coconut endocarp fractions were separated primarily to enable selection of samples for AMS dating.

When all of the test units are compared directly, the highest quantities of coconut endocarp were observed within Talasiu TP2 and Heketa TP3 with a total of 1.6g in each test unit. However, when these quantities are considered as a fraction of the total macrobotanical weight for each test unit, the highest percentages were extracted from Leka TP3 (10.7%) and TP4 (19.5%) (see Table 8.2). The remaining test units contained coconut endocarp fractions that 139 were less than 10% of the entire macrobotanical weight, with Leka TP3 containing the smallest percentage.

Recovered coconut endocarp was distributed throughout the test unit from Talasiu, with quantities found in every stratigraphic layer, aside from the sterile clay at the base, and almost every spit. Only one stratigraphic layer was processed in TP2 from Leka. This layer was Layer 8, which was a mid-brown silty clay cultural layer with charcoal, and this contained coconut endocarp in the bottom 20cm (105-125cm below surface) before sterile clay was encountered. Leka TP3 only contained coconut endocarp in this same cultural layer in the top 5cm (90- 95cmbs). In contrast, coconut endocarp was extracted from all sampled stratigraphic layers in Leka TP4 (Layers 4-6), but not every spit in these. Small quantities were recovered from the lower 15cm (110-125cmbs) of Layer 4, a mid-brown silty clay layer similar to that observed in TP2 and TP3, and the top 5cm (125-130cmbs) and middle (134-140cmbs) of Layer 5 in a more dense shell deposit. Coconut endocarp was also recovered from the bottom 10cm (155- 170cmbs) of this test unit in the mixed clay and shell deposit of Layer 6. At Heketa, coconut was extracted from all stratigraphic layers in TP2, but was not observed within the bottom 10cm of this test unit (90-100cmbs). Coconut endocarp extracted from TP3 was mostly concentrated in the lower 35cm (70-105cmbs) of this test unit, in the base of Layer 3 and throughout Layers 4-5.

Site TP Wet-sieved (g) In-situ (g) Flot (g) Total % of total macrobotanical weight Talasiu 2 1.6 0 0 1.6 2.6 Leka 2 0.6 0 0 0.6 6.4 Leka 3 0.1 0 0 0.1 2.1 Leka 4 0.7 0 0 0.7 10.7 Heketa 2 1.6 0 0 1.6 19.5 Heketa 3 1.2 0 0 1.2 4.0 Total 5.8 0 0 5.8 45.3

Table 8.2 Quantification of coconut endocarp from all sites and test-pits

Other endocarp Both wood charcoal and non-coconut endocarp composed a large percentage of the macrobotanical remains from all sites and test units. Endocarp was successfully extracted using all three archaeobotanical techniques employed during fieldwork. Wet-sieving and in situ collection resulted in the extraction of endocarp from all test-units, whilst flotation had more variable rates of recovery. This could be the result of issues with the flotation of endocarp already highlighted with regard to coconut endocarp extraction. This fraction was separated as a preliminary step towards the selection of samples for AMS dating. These endocarp fractions remain unidentified because of the lack of a comprehensive comparative collection. Some Aleurites moluccana was observed but was not separated from other endocarp due to a focus within this study on the identification of parenchyma within the macrobotanical remains.

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The quantification of other endocarp followed the same pattern as coconut endocarp, whereby the highest quantities were extracted from Talasiu TP2 (5.8g) and Heketa TP3 (5.1g) (see Table 8.3). When quantities of endocarp are considered as percentages of the total weight of macrobotanical material extracted from these test units, Leka TP4 clearly has the highest percentage of other endocarp (62.9%), while Leka TP2 has the lowest (3.2%) by a significant margin.

Talasiu test unit TP2 again has recovered endocarp distributed throughout all strata. Only two spits did not contain any non-coconut endocarp, at 0-5cmbs and 80-85cmbs. The three test units at Leka have variable distributions of endocarp throughout the observed stratigraphic layers. Endocarp was recovered from the top (90-95cmbs), middle (105-110cmbs) and bottom (120-125cmbs) spits of Layer 5 in TP2. In contrast, TP3 had non-coconut endocarp in all three sampled spits of the same stratigraphic layer, while TP4 had small quantities of endocarp throughout the test unit apart from the top 5cm (95-100cmbs). Non-coconut endocarp was found throughout all strata within TP2 and TP3 at Heketa, but was absent from the bottom 5cm (100- 105cmbs) of TP3.

Spit TP Wet-sieved (g) In-situ (g) Flot (g) Total % of total macrobotanical weight Talasiu 2 3.6 0.9 1.3 5.8 9.5 Leka 2 0.2 0.1 0 0.3 3.2 Leka 3 0.6 1.4 0.3 2.3 47.9 Leka 4 2.4 1.7 0 4.1 62.9 Heketa 2 1.6 0.3 0.7 2.6 31.7 Heketa 3 3.9 0.7 0.5 5.1 16.8 Total 12.3 5.1 2.8 20.2 172.0

Table 8.3 Quantification of other endocarp from all test units

Wood charcoal and parenchyma The separation of monocot and dicot wood and parenchymatous charcoal from coconut and other endocarp was the final stage in the quantification of macrobotanical material. Because of the similarity of vegetative storage parenchyma to wood charcoal, any plant-derived charred material that was not endocarp was separated first and weighed as a single fraction. Because of the abundance of charcoal from Talasiu TP2, this unit was used as a test for identifiable parenchymatous material. Wet-sieving and flotation techniques were by far the most successful method for recovering both wood charcoal and parenchyma. The vesicular cellular structure and specific gravity of wood charcoal enables this material to float relatively easily in water, a fact which has been utilised by archaeobotanists for many years. The success of wet-sieving, as a follow-up to flotation steps, is most likely the result of the inability of technicians to easily deflocculate clay sediments to release wood charcoal in the field using bucket flotation methods.

The highest quantities of wood charcoal (including parenchyma) were recovered from test unit TP2 at Talasiu with a total of 53.4g (see Table 8.4). TP3 at Heketa had the second greatest quantity (24.1g) of wood charcoal, while the remaining test units all had less than 10g. 141

Despite these figures, when these quantities are viewed as percentages of the whole macrobotanical assemblages from each of these test units, Leka TP2 has the highest percentage of wood charcoal, with 90.4%. These statistics indicate that the assemblages from Talasiu TP2, Leka TP2 and TP3, and Heketa TP3 are all dominated by wood charcoal. In contrast only 26.4% of recovered charcoal is represented by wood charcoal in Leka TP4, while Heketa TP3 has just under half (48.8%).

The distribution of these quantities within the test units from the sites of Talasiu, Leka and Heketa are variable; however, at least small amounts of wood charcoal were present in every spit and stratigraphic layer sampled. In Talasiu TP2, most charcoal was concentrated around Layer 3 (40-75cm) where it is likely that larger fragments were preserved in the matrix of loose shell and yellow red clay. Very little charcoal was recovered from Layers 1, 2 and 5 in the more compact shell midden deposits. Leka TP2, TP3 and TP4, and Heketa TP2 had relatively even distribution of wood charcoal in the test units, with small amounts (<2g) in every spit and stratigraphic layer. In contrast, Heketa TP3 had the largest concentration of charcoal within Layer 4 (80-90cmbs) with 12.5g, while the remaining strata had less than 5g.

Spit TP Wet-sieved (g) In-situ (g) Flot (g) Total % of total macrobotanical weight Talasiu 2 23.7 17.9 11.8 53.4 87.8 Leka 2 3.5 4.5 0.5 8.5 90.4 Leka 3 0.8 1.6 0.0 2.4 50.0 Leka 4 1.0 0.3 0.5 1.7 26.4 Heketa 2 2.0 1.1 0.9 4.0 48.8 Heketa 3 22.4 1.2 0.5 24.1 79.3 Total 53.4 26.6 14.1 94.0 382.6

Table 8.4 Quantification of wood charcoal and parenchyma from all test units

Parenchyma distribution and identification: Talasiu TP2 case study Within this section the macrobotanical assemblage from test unit TP2 at Talasiu (TO-Mu-2) was chosen to conduct a case study for the separation and identification of vegetative storage parenchyma. Due to the documented difficulty of separating these charred remains from wood charcoal (Hather 2000), it was decided to analyse only one of the six test units excavated during the 2011 field season. Talasiu TP2 was chosen for the abundance of charcoal in the macrobotanical assemblage of this unit, on the assumption that the preservation of any vegetative storage parenchyma was likely to be the greatest.

Parenchyma was first separated from wood charcoal based on ground tissue morphology, such as consistent cell shapes that are usually rounded or angular; the presence of distinctive vascular bundles or tissues, usually with very few rays dividing these tissues. This initial step enabled the presence of vegetative and non-vegetative parenchyma (i.e. fruits) in samples to be recorded. The distribution of these remains in the test unit varied, although at least one fragment of charred parenchyma was extracted from each stratigraphic layer. The disturbed

142 midden within Layer 1, and very compact shell midden deposit in Layer 2 both contained three fragments. The largest overall quantity of parenchyma was recovered from the loose large shell matrix composing Layer 3 (see Table 8.5), with a total of five fragments. Layer 4 below this contained another two fragments, while the basal cultural layer of reddish clay and small crushed shell only contained one fragment.

Each fragment was identified to species where possible, using a range of attributes of ground tissue and vascular tissue morphology, and the identification flowchart key created for the reference collection (Chapter 5). Most fragments could be identified with a moderate degree of confidence, and were therefore labelled with the prefix ‘cf.’, indicating that most morphological features matched the written descriptions and resembled the reference SEM images.

A number of different families are represented within the identified parenchyma assemblage. These included Araceae, Dioscoreaceae, Moraceae, Musaceae, and Zingiberaceae. Four fragments were identified as cf. Colocasia esculenta, or the common taro and were extracted from Layers 2 and 3 to a maximum depth of 65cmbs. Two species belonging to the Dioscorea or yam genus, Dioscorea alata and Dioscorea nummularia, were found in Layers 3 and 4, with a total of four identified fragments. A single fragment identified as cf. Artocarpus altilis (breadfruit) fruit/flesh was found in Layer 2 between 25-30cmbs. Interestingly, a single fragment of parenchyma from Layer 3 was identified as belonging to the Musaceae family, indicating that the flesh of this fruit was either intentionally cooked or discarded into a fire and incorporated into the ash. Members of the Ginger family Zingiberaceae, such as Zingiber zerumbet or Curcuma longa were often cultivated and eaten in Tonga or used for medicinal and ornamental purposes. The antiquity of this use is demonstrated by the recovery of parenchyma identified confidently to this family within the archaeobotanical record at Talasiu. The remaining fragments could only be identified as ‘root-derived’ (Layer 3), or were left classified as ‘unidentifiable’ (Layers 1 and 5).

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Depth Spit Layer Parenchyma Identification 0-5 1 1 5-10 2 1 Unidentifiable 10-15 3 1 x Zingiberaceae 15-20 4 1 x Zingiberaceae 20-25 5 2 x cf. Colocasia esculenta 25-30 6 2 x cf. Artocarpus altilis fruit 30-35 7 2 x cf. Colocasia esculenta 35-40 8 3 cf. Dioscorea alata 40-45 9 3 x cf. Musaceae 45-50 10 3 x cf. Colocasia esculenta 50-55 11 3 x cf. Dioscorea alata 55-60 12 3 x Root-derived 60-65 13 3 x cf. Colocasia esculenta 65-70 14 4 70-75 15 4 x cf. Dioscorea alata 75-80 16 4 x cf. Dioscorea nummularia 80-85 17 5 85-90 18 5 x Unidentifiable 90-95 19 5 95-100 Control 6

Table 8.5 Distribution and identification of parenchyma extracted from Talasiu TP2 Microbotanical analysis

Extraction, quantification and distribution Microbotanical assemblage breakdown Starch was successfully extracted from all test units sampled for microbotanical remains from Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2). Due to time constraints, only four of the six excavated test units were sampled, with a total of 39 bulk soil samples processed. These test units included Talasiu TP2, Leka TP2 and TP4, and Heketa TP3, and were chosen based on site taphonomy and the likelihood of microbotanical preservation. Of these, Talasiu TP2 contained the largest quantity (count) of starch grains, but this is biased by the large number of samples analysed from this site. Every spit or level was sampled from Talasiu TP2, whilst every second spit was sampled from the remaining test units. This alteration of the microbotanical subsampling procedure was made after Talasiu TP2 was completely processed, and counts revealed that there was very little variation in starch distribution within the test unit apart from a spike at the very base of the cultural deposits. Due to time constraints, it was deemed appropriate to process every second sample and halve the overall processing time.

With this in mind, it may be appropriate to compare the quantities of starch from each of the test units as an average based on the number of processed samples (see Table 8.6). When these figures are calculated, it becomes clear that Leka TP2 had the highest quantity of starch per sample, with an average of 93 grains. Leka TP3 followed with an average of 31.4 grains per

144 sample, just slightly higher than Talasiu TP2 which contained an average of 22.5 grains per sample. The lowest average derived from TP3 at Heketa (9 grains). Considering that AMS dating indicates that this site was the most recently occupied, these figures would indicate that either plant food was not being processed or discarded in these cultural deposits at Heketa, or that taphonomic factors such as soil pH levels or enzymatic activity heavily impacted starch preservation. These quantities from each test unit will be briefly discussed with regard to the distribution of starch throughout the sampled cultural deposits below.

Site Total samples analysed Total count Average/sample Talasiu TP2 20 449 22.5 Leka TP2 4 372 93.0 Leka TP4 8 251 31.4 Heketa TP3 7 63 9.0 Totals 39 1135 29.1

Table 8.6 Overall quantities (counts) of starch extracted from all sampled test units at Talasiu (TO-Mu-2)

Talasiu (TO-Mu-2) Test unit TP2 from Talasiu contained a total of 449 starch grains, extracted from 20 processed 3gm bulk soil samples (see Table 8.7). When this quantity is broken down into the distribution of starch grains within each recorded stratigraphic layer, some patterning emerges. The disturbed midden at the top of the test unit (0-20cmbs) which composes Layer 1 has a total of 107 starch grains from four samples. Layer 2 below this has the smallest quantity of starch grains, with a total of 47 grains from three samples. Layer 3 has the largest number of samples (n=6), however this deposit has a quantity of starch (96 grains) that falls in the middle of the range when compared to the other strata. Layer 4 had the second smallest quantity of starch with 64 grains from three samples. Finally, Layer 5 at the base of the cultural deposits had the largest quantity, with a total of 128 grains extracted from three samples. The control sample taken from the sterile clay below the shell midden had seven starch grains, which most likely derived from the transition between these two deposits.

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Depth Spit Layer Starch count 0-5 1 1 11 5-10 2 1 19 10-15 3 1 34 15-20 4 1 43 20-25 5 2 11 25-30 6 2 11 30-35 7 2 25 35-40 8 3 14 40-45 9 3 5 45-50 10 3 20 50-55 11 3 6 55-60 12 3 46 60-65 13 3 5 65-70 14 4 16 70-75 15 4 10 75-80 16 4 38 80-85 17 5 18 85-90 18 5 24 90-95 19 5 86 95-100 Control 6 7 Totals 449

Table 8.7 Distribution of starch counts within Talasiu TP2

Leka (J17) Two test units were sampled from excavations at Leka, TP2 and TP4. Leka TP3 was excluded from this analysis as stratigraphic comparisons revealed that the cultural deposit sampled within this test unit was already sampled in TP2. TP2 had a total starch count of 372 starch grains extracted from four 3gm bulk soil samples. These were distributed relatively evenly throughout the test unit, which seems reasonable considering these four samples were taken from the same stratigraphic layer (Layer 7) (see Table 8.8). Slightly fewer starch grains were recovered in the lower 10cm of the unit, above the sterile clay. This is most likely a result of sampling as these figures are not significantly different from the samples above, especially when considered in comparison with other samples from other test units.

Depth Spit Layer Starch count 90-95 1 7 102 100-105 3 7 100 110-115 5 7 82 120-125 7 7 88 Totals 372

Table 8.8 Distribution of starch counts within Leka TP2

Leka TP4 had a different starch distribution pattern. Most starch was located in Layer 4 in two samples from 95-110cmbs, where the 100 grain maximum count was reached for both samples. Layer 4 is possibly the same as Layer 7 in TP2, but the distance between these two units makes this correlation difficult to prove. However, this would explain the high quantity of

146 starch extracted from these deposits, especially in comparison with other deposits in TP4. Below Layer 4, starch quantities drop off significantly in Layers 5-6. Layer 5 has a total of 30 starch grains from three samples, while Layer 6 only has five grains from two samples.

Depth Spit Layer Starch count 95-100 1 4 100 105-110 3 4 100 115-120 5 4 16 125-130 7 5 9 135-140 9 5 16 145-150 11 5 5 155-160 13 6 5 165-170 15 6 0 Totals 251

Table 8.9 Distribution of starch counts within Leka TP4

Heketa (TO-Nt-2) Only one test unit from Heketa was sampled for microbotanical remains. A decision was made to sample TP3, and leave TP2 due to both time constraints, and the fact that stratigraphic comparison indicated that the deposit sampled in TP2 (Layer 4) was most likely the same as Layer 3 in TP3. As mentioned earlier, TP3 had the smallest amount of extracted starch with a total of 63 starch grains from seven samples. Similar to Leka TP3, the highest quantity of starch was extracted from the uppermost layer, Layer 3, with 43 grains recorded (see Table 8.10). Below this, Layer 4 had 11 grains extracted from two samples, while Layer 5 had only seven grains. Layer 5 is an ash lens within Layer 4, and this could explain the low quantity in this sample.

Depth Spit Layer Starch count 40-45 1 3 25 50-55 3 3 9 60-65 5 3 2 70-75 7 3 7 80-85 9 4 9 90-95 11 5 7 100-105 13 4 4 Totals 63

Table 8.10 Distribution of starch counts within Heketa TP3

Identification: Assemblage-typology approach As Talasiu TP2 was the first test unit to be processed for starch and classified using multivariate statistics in the form of Linear Discriminant Function Analysis, this unit was utilised as a case study. The assemblage-typology approach was compared with the DFA classifications, followed by visual checking of starch grains. All of the extracted archaeological starch was recorded using the same morphological attributes as the comparative collection and entered into an Excel 147 database. Once this was complete, it was important to establish what morphological patterning could be seen within the assemblage. Therefore, a number of starch types were created that were based on a combination of the recorded nominal attributes. The primary variable used was shape, and then further attributes such as the number and type of pressure faceting, and the presence and type of fissuring at the hilum. The abundance of these starch types in the assemblage from Talasiu TP2 was calculated. However, as we still do not fully understand factors affecting starch production with plant organs and starch preservation in sediments, quantification may not be the best analytical technique to use here. Simple presence or absence was deemed to be more appropriate and was used for further analysis.

In order to establish the identity of starch types they were compared to those grains in the comparative collection. Most often, a number of possible matches were made. To narrow this down further, the archaeological starch types were compared to the length range of the matches. For example, Type 1 or cone-shaped grains, were present in the yams Dioscorea alata, Dioscorea bulbifera, Dioscorea rotundata and Dioscorea nummularia. It is possible to rule out D. rotundata based on length, and probably also D. nummularia (see Figure 8.2). Therefore this starch type most likely originated from D. alata or D. bulbifera, but it is probably most appropriate to keep this identification at family level at this stage (see Table 8.11).

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Figure 8.2 Box plot demonstrating maximum length comparison of archaeological starch Type 1 with Dioscorea spp.

Table 8.11 Table outlining suggested family of origin of archaeological starch types from Talasiu TP2.

This preliminary starch identification suggests that the Talasiu assemblage contains starch from Dioscoreaceae, Musaceae and from the Araceae or aroid family, in this case most likely originates from Cyrtosperma merkusii or the giant swamp taro, and Amorphophallus paeoniifolius, the giant stink lily, rather than Colocasia esculenta or the common taro. Interestingly, comparison of these archaeological starch grains with reference specimens not

149 expected to be present within the time period represented by the deposits at Talasiu tentatively identified some of the starch as possibly originating from the Convolvulaceae family to which Ipomoea batatas or sweet potato belongs. Clearly, this species should not be in the assemblage as I. batatas is thought to be a late prehistoric or historic introduction into Tonga (Fall 2010; Kirch 1978, 1990), and so is either contamination or a misidentification. Later multivariate statistics and visual checking ruled out the identification of I. batatas for Talasiu TP2. The distribution of identifications suggested by the assemblage- typology approach is outlined in Table 8.12.

The assemblage-typology approach suggests some basic family-level identifications of archaeological starch from Talasiu TP2. However, the use of multivariate statistics in the form of Discriminant Function Analysis enabled these identifications to be more confidently narrowed to species for a larger number of extracted grains. In light of the more accurate species-level data provided by multivariate statistical analysis, assemblage-typology analysis was not applied to the rest of the sites and test units from Leka (J17) and Heketa (TO-Nt-2). For the multivariate statistics, a comprehensive reference collection was available containing both nominal and metric data with light microscope and SEM imagery, enabling the comparison of reference material data with that from extracted archaeological starch.

Layer Spit Depth Dioscorea sp. Musaceae Araceae 1 1 0-5cm • 1 2 5-10cm • 1 3 10-15cm • 1 4 15-20cm • • 2 5 20-25cm • • 2 6 25-30cm • 2 7 30-35cm • • 3 8 35-40cm • 3 9 40-45cm 3 10 45-50cm • • 3 11 50-55cm 3 12 55-60cm • 3 13 60-65cm 4 14 65-70cm • 4 15 70-75cm • 4 16 75-80cm • 5 17 80-85cm • 5 18 85-90cm • • 5 19 90-95cm • • • 6 Control 95-100cm

Table 8.12 Distribution of preliminary identifications within Talasiu TP2 using the assemblage-typology approach

Identification: Multivariate statistics—Discriminant Function Analysis Linear Discriminant Function Analysis (DFA) within the PAST statistical software package was utilised as part of a multivariate statistical approach to the identification of archaeological starch

150 grains. The methods for applying this technique within the reference collection to assess the morphological diversity of Pacific cultigens (Chapters 3-5), and how this statistical analysis would enable the classification of archaeological starch with varying degrees of confidence (Chapter 8) have already been explained. The results of this analysis will be presented here, along with the steps taken to create a species list of high confidence for each of the three sites under analysis.

Starch classifications were made using the same algorithms developed for the reference collection, and were assessed with varying degrees of confidence based on the species-level data. In order to interpret archaeological classifications each sample was analysed according to the number of starch grains classified to each species, and the percentage of correct reclassifications of that species when the reference collection comparison stage had been carried out (Chapter 5). To reiterate, these criteria were:

 High confidence classifications had to have a successful reclassification rate within DFA for that species of over 60%, and more than five grains had been matched to that species within that sample.  Moderate confidence classifications had to either have over 60% correct reclassification but less than five grains matched, or less than 60% correct reclassification and over five grains matched to that species within that sample.  Low confidence classifications were given when the reclassification for that species was less than 60%, and less than five grains were matched to that species.

The database of archaeological starch extracted from each sample (spit) was divided into two based on the orientation of each starch grain (eccentric/side on or centric/end-on). These were compared to the two datasets created for the reference collection. The outputs of the DFA produced a list of classifications, as well as Discriminant Analysis plots showing the distribution of archaeological grains according to the first two canonical variates (see Figure 8.3 and Figure 8.4). These act as visual guides for inspection of Mahalanobis distances from group centroids and ellipses (Baxter 2003) and Figure 8.3suggests that many of the archaeological grains classified in the centric dataset may in fact belong to none of these groups. In contrast, in Figure 8.4 the distribution of archaeological grains within the eccentric reference dataset more often falls within the ellipses of these species, in some species more commonly than others.

Talasiu (TO-Mu-2) All of the 449 starch grains extracted from 3g soil sub-samples from TP2 at Talasiu were assigned to a species using DFA. When these classifications were tabulated according to sample (spit) and level of confidence, it became clear that the archaeological starch grains were assigned to most of the species within the reference collection with at least low level confidence. The only species not apparently represented by archaeological starch were

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Xanthosoma sagittifolium (modern taro introduction), Curcuma longa (turmeric) and Dioscorea pentaphylla (five-leaved yam). Several species could be classified with high confidence within the test unit. These included Colocasia esculenta (common taro), Inocarpus fagifer (Tahitian chestnut), Musa sp.2 (banana), Piper methysticum (kava) although this could be considered a contaminant, and Spondias dulcis (Otaheite apple). Some starch was also classified with low level confidence to suspected historic or modern contaminants, including Ipomoea batatas (sweet potato), which was probably a late prehistoric introduction and Solanum tuberosum (white/Irish potato), which is a modern contaminant. These present problems for the dataset, but visual checking of the starch classifications within the next stage of the analysis proved successful in eliminating misclassifications from the final species lists.

Figure 8.3 Discriminant analysis plot for centric dataset showing ellipses (coloured dots represent reference species, black dots represent archaeological grains)

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Figure 8.4 Discriminant analysis plot for eccentric dataset showing ellipses (coloured dots represent reference species, black dots represent archaeological grains)

Depth Layer Spit macrorrhiza Alocasia Aroid Taro) (Giant paeoniifolius Amorphophallus Foot (Elephant Yam) Aroid Artocarpus altilis (Breadfruit) asiatica Barringtonia Tree) Poison (Fish esculenta Colocasia (Common Aroid Taro) Curcuma longa (Turmeric) Cyrtosperma merkusii Aroid Swamp Taro) (Giant alata Dioscorea (Common Yam) bulbifera Dioscorea (Bitter Yam) esculenta Dioscorea (Lesser Yam) nummularia Dioscorea Yam) (Spiny fagifer Inocarpus chestnut) (Tahitian batatas Ipomoea (Sweet Potato) CONTAMINATION citrifolia Morinda mulberry)(Indian sp. 1 Musa (Banana) sp. 2 Musa (Banana) Piper methysticum (Kava) Pteridium sp. (Bracken) tuberosumSolanum (White Potato) CONTAMINATION dulcis Spondias (Otaheite Apple or Ambarella) leontopetaloides Tacca Arrowroot)(Polynesian sagittifolium Xanthosoma (Arrowleaf Taro) Ear Elephant Aroid CONTAMINATION 0-5 1 1 5-10 1 2 10-15 1 3 15-20 1 4 20-25 2 5 25-30 2 6 30-35 2 7 35-40 3 8 40-45 3 9 45-50 3 10 50-55 3 11 55-60 3 12 60-65 3 13 65-70 4 14 70-75 4 15 75-80 4 16 80-85 5 17 85-90 5 18 90-95 5 19 95-100 6 Control

Table 8.13 Levels of confidence from DFA classification of archaeological starch from Talasiu TP2. NB High confidence (black), moderated confidence (medium grey) and low confidence (light grey)

Using visual checking techniques, many classifications were rejected, leaving a much smaller final species list for Talasiu TP2. Some obvious misclassifications were also able to be corrected using this step, and new classifications provided. Three aroids were represented after visual assessment in this final list, including Amorphophallus paeoniifolius (elephant foot yam), Colocasia esculenta, and Cyrtosperma merkusii (giant swamp taro), along with two yams— Dioscorea alata and Dioscorea bulbifera, two bananas, Piper methysticum (kava), and several arboreal species including Artocarpus altilis (breadfruit), Barringtonia asiatica (fish poison 153 tree), Inocarpus fagifer, and Spondias dulcis. These were tabulated according to the presence of these species, as quantification of classifications at this final stage was not made due to starch preservation and taphonomy. The distribution of these species in the test unit may also be subject to these same issues of taphonomy and preservation.

Despite this, some general patterning noted during analysis will be discussed here. Of particular interest was the observation that several species were present throughout most stratigraphic layers within the test unit, notably A. paeoniifolius and I. fagifer present in every layer. These are not primary crops, but are instead recorded ethnographically and through oral traditions as supplementary or famine foods. Only one species, Musa sp. 1, was present in only a single deposit (Layer 5). Some interesting stratigraphic patterning emerges in the range of species recorded within each deposit. Layer 1 (0-20cmbs) contains starch from all three aroids, both yams, I. fagifer, P. methysticum and S. dulcis. Layer 2 (20-35cmbs) also contains starch from three aroids, but only one of the yams— D. alata, along with several arboreal species including B. asiatica, I. fagifer and S. dulcis. Layer 3 is the largest (in terms of thickness) of all of the deposits recorded within TP2, and has the largest number of species, including all the previously mentioned species apart from D. bulbifera and Musa sp.1. Layer 4 contains two of the aroids, A. paeoniifolius and C. merkusii, a single yam species— D. alata, P. methysticum, and a number of arboreal species including A. altilis, B. asiatica, and I. fagifer. The lowermost cultural deposit, Layer 5, contained a fairly comprehensive species list, which included all three aroids, D. bulbifera, both Musa sp., A. altilis, B. asiatica, I. fagifer, P. methysticum and S. dulcis.

Depth Layer Spit paeoniifolius Amorphophallus Aroid Yam) Foot (Elephant altilis Artocarpus (Breadfruit) asiatica Barringtonia Tree)(FishPoison esculenta Colocasia Aroid Taro) (Common merkusii Cyrtosperma Aroid Taro) Swamp (Giant alata Dioscorea Yam) (Common bulbifera Dioscorea Yam) (Bitter fagifer Inocarpus chestnut) (Tahitian 1 sp. Musa (Banana) 2 sp. Musa (Banana) methysticum Piper (Kava) dulcis Spondias Ambarella) Apple(Otaheiteor 0-5 1 1 5-10 1 2 10-15 1 3 15-20 1 4 20-25 2 5 25-30 2 6 30-35 2 7 35-40 3 8 40-45 3 9 45-50 3 10 50-55 3 11 55-60 3 12 60-65 3 13 65-70 4 14 70-75 4 15 75-80 4 16 80-85 5 17 85-90 5 18 90-95 5 19 95-100 6 Control

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Table 8.14 Final table documenting species represented by archaeological starch within Talasiu TP2 NB Presence indicated by black squares

Leka (J17) In total, 623 starch grains were extracted from 12 bulk soil samples taken from the two excavated test units at Leka (J17), and each of these grains were again classified using DFA to a suggested species based on the statistical data collected for the reference collection. Each of these two test units will be analysed separately first, and then the two sets of final classifications will be discussed together.

The first of these units, TP2, had four soil samples processed for starch extraction. From these samples, a large list of possible classifications was produced using DFA, and interpreted with varying levels of confidence. As with the classifications provided for Talasiu TP2, most species in the reference collection were represented in the list of assignments for the archaeological starch from Leka TP2. Only four species- Alocasia macrorrhiza, Curcuma longa, Morinda citrifolia (Indian mulberry), and Pteridium sp. (bracken fern) were not included in the preliminary list of assignments. Also similar to Talasiu TP2, several of the same species at Leka TP2 were often classified with high confidence throughout the test unit, including A. paeoniifolius, I. fagifer and S. dulcis. This is most likely a direct result of the high reclassification rate of these species within the multivariate analysis of starch morphological attributes within the reference collection, which created the algorithm for the classification of archaeological starch. Many other species were classified with moderate confidence, and very few with low confidence. Some potential contaminants were identified within this test unit, including I. batatas and S. tuberosum distributed throughout the unit, with low or moderate confidence.

Depth Layer Spit paeonniifolius Amorphophallus Aroid Yam) Foot (Elephant altilis Artocarpus (Breadfruit) asiatica Barringtonia Tree)(FishPoison esculenta Colocasia Aroid Taro) (Common merkusii Cyrtosperma Aroid Taro) Swamp (Giant alata Dioscorea Yam) (Common bulbifera Dioscorea Yam) (Bitter esculenta Dioscorea (LesserYam) nummularia Dioscorea Yam) (Spiny fagifer Inocarpus chestnut) (Tahitian batatas Ipomoea CONTAMINATION (SweetPotato) 1 sp. Musa (Banana) 2 sp. Musa (Banana) methysticum Piper (Kava) tuberosum Solanum CONTAMINATION (WhitePotato) dulcis Spondias Ambarella) Apple(Otaheiteor leontopetaloides Tacca (PolynesianArrowroot) 90-95 7 1 100-105 7 3 110-115 7 5 120-125 7 7

Table 8.15 Levels of confidence from DFA classification of archaeological starch from Leka TP2. NB High confidence (black), moderated confidence (medium grey) and low confidence (light grey)

Once these classifications were assessed using visual checking, a final smaller list of species was made for the archaeological starch from Leka TP2. These species were all from the same cultural deposit (Layer 7), as this was the only stratigraphic layer sampled from this test unit, but there is some variation in the distribution of these species at different depths within this layer. Several species are present throughout the deposit, including Amorphophallus

155 paeoniifolius, Cyrtosperma merkusii and Inocarpus fagifer. Artocarpus altilis is identified in all spits apart from Spit 5 (110-115cmbs), while Spondias dulcis is present in all but Spit 1 (90- 95cmbs). A number of species are present in only one sample, including Colocasia esculenta, Barringtonia asiatica, Curcuma longa (identified only using visual checking), Dioscorea esculenta, Musa sp.1, and Piper methysticum. The significance of these identifications will be discussed later in this section.

Depth Layer Spit paeoniifolius Amorphophallus Aroid Yam) Foot (Elephant altilis Artocarpus (Breadfruit) asiatica Barringtonia Tree) (Fish Poison esculenta Colocasia Aroid Taro) (Common longa Curcuma (Turmeric) merkusii Cyrtosperma Aroid Taro) Swamp (Giant esculenta Dioscorea Yam) (Lesser fagifer Inocarpus chestnut) (Tahitian 1 sp. Musa (Banana) methysticum Piper (Kava) dulcis Spondias Ambarella) or Apple(Otaheite 90-95 7 1 100-105 7 3 110-115 7 5 120-125 7 7

Table 8.16 Final table documenting species represented by archaeological starch within Leka TP2

A larger number of samples were processed for Leka TP4 (n=7), although this did not result in greater quantities of extracted starch. The preliminary classifications of this starch produced a list of low-to-high confidence assignments that was quite different to that of TP2, although again four species from the reference collection were not included in this list— Alocasia macrorrhiza, Xanthosoma sagittifolium, Curcuma longa and Dioscorea pentaphylla. Very few species were classified with high confidence in any sample. Those species that were classified with high confidence included Barringtonia asiatica, Inocarpus fagifer, and Musa sp. 2, and represented a mix of cultivated and semi-cultivated species. A number of other species could primarily be classified with only low level confidence, including Cyrtosperma merkusii, Artocarpus altilis, Dioscorea esculenta, Dioscorea nummularia, Ipomoea batatas and Tacca leontopetaloides. The remaining species were mostly classified with moderate level confidence due to low numbers or low re-classification rates in the reference collection.

Depth Layer Spit paeonniifolius Amorphophallus Aroid Yam) Foot (Elephant altilis Artocarpus (Breadfruit) asiatica Barringtonia Tree)(FishPoison esculenta Colocasia Aroid Taro) (Common merkusii Cyrtosperma Aroid Taro) Swamp (Giant alata Dioscorea Yam) (Common bulbifera Dioscorea Yam) (Bitter esculenta Dioscorea (LesserYam) nummularia Dioscorea Yam) (Spiny fagifer Inocarpus chestnut) (Tahitian batatas Ipomoea CONTAMINATION (SweetPotato) citrifolia Morinda mulberry) (Indian 1 sp. Musa (Banana) 2 sp. Musa (Banana) methysticum Piper (Kava) sp. Pteridium (Bracken) tuberosum Solanum CONTAMINATION (WhitePotato) dulcis Spondias Ambarella) Apple(Otaheiteor leontopetaloides Tacca (PolynesianArrowroot) 95-100 4 1 105-110 4 3 115-120 4 5 125-130 5 7 135-140 5 9 145-150 5 11 155-160 6 13

Table 8.17 Levels of confidence from DFA classification of archaeological starch from Leka TP4. NB High confidence (black), moderated confidence (medium grey) and low confidence (light grey) 156

The final analysis of these classifications produced a list of species containing a range of both primary cultivated and supplementary species. As observed in TP2, the presence of Solanum tuberosum indicates some contamination within these samples (likely from the laboratory environment). Layer 4 (95-120cmbs) contained the largest number of species to which the extracted archaeological starch could be confidently assigned. From within this layer of mixed shell and clay matrix, starch deriving from three aroids— Amorphophallus paeoniifolius, Cyrtosperma merkusii and Colocasia esculenta, four yams— Dioscorea alata, Dioscorea bulbifera, Dioscorea esculenta and Dioscorea nummularia, and a number of tree crops and supplementary species such as Artocarpus altilis, Barringtonia asiatica, Inocarpus fagifer, one Musa sp. and Spondias dulcis were extracted. Layer 5 (125-150cmbs) had a much smaller list of species present, including one aroid— C. merkusii, one yam— D. alata, P. methysticum (which again may also be a contaminant and will be discussed in Chapter 9), S. tuberosum, and arboreal species such as A. altilis, I. fagifer and S. dulcis. Layer 6 (155- 160cmbs) had only one species that could be confirmed using visual checking, being S. dulcis.

Depth Layer Spit paeoniifolius Amorphophallus Aroid Yam) Foot (Elephant altilis Artocarpus (Breadfruit) asiatica Barringtonia Tree)(FishPoison esculenta Colocasia Aroid Taro) (Common merkusii Cyrtosperma Aroid Taro) Swamp (Giant alata Dioscorea Yam) (Common bulbifera Dioscorea Yam) (Bitter esculenta Dioscorea (LesserYam) nummularia Dioscorea Yam) (Spiny fagifer Inocarpus chestnut) (Tahitian 2 sp. Musa (Banana) methysticum Piper (Kava) tuberosum Solanum CONTAMINATION (White Potato) dulcis Spondias Ambarella) Apple(Otaheiteor 95-100 4 1 105-110 4 3 115-120 4 5 125-130 5 7 135-140 5 9 145-150 5 11 155-160 6 13

Table 8.18 Final table documenting species represented by archaeological starch within Leka TP4

When the classifications from these two test units from Leka (J17) are combined, a much larger overall list of species identified as being utilised from 1300 to 1000 cal BP. This list includes three aroids— A. paeoniifolius, C. merkusii and C. esculenta, four yams— D. alata, D. bulbifera, D. esculenta and D. nummularia, P. methysticum, a number of tree crops and supplementary species such as A. altilis, B. asiatica, I. fagifer, two Musa spp., and S. dulcis.

Heketa (TO-Nt-2) The single test unit processed for starch at Heketa, TP3, contained a total of 63 starch grains extracted from seven samples, indicating a very low level of starch preservation compared to that observed within all of the other test units. Despite this, a number of low-to-high level confidence classifications were assigned to these grains using DFA. Only two species could be assigned to starch with high confidence in any of the samples, and these were Inocarpus fagifer and Spondias dulcis within Spit 1 (40-45cmbs) from Layer 3. These were also classified with 157 moderate confidence in other samples. Another seven species were identified with predominantly moderate levels of confidence, and this list includes Amorphophallus paeoniifolius, Barringtonia asiatica, Dioscorea bulbifera, Musa sp. 2, Piper methysticum, Pteridium sp., and Solanum tuberosum (contaminant). Six remaining species could only be classified with low confidence, including Alocasia macrorrhiza, Cyrtosperma merkusii, Artocarpus altilis, Dioscorea esculenta, and Ipomoea batatas. The remaining species from the reference collection were not represented by any archaeological starch from TP3.

Depth Layer Spit macrorrhiza Alocasia Aroid Taro) (Giant paeoniifolius Amorphophallus Aroid Yam) Foot (Elephant altilis Artocarpus (Breadfruit) asiatica Barringtonia Tree)(FishPoison merkusii Cyrtosperma Aroid Taro) Swamp (Giant bulbifera Dioscorea Yam) (Bitter esculenta Dioscorea (LesserYam) fagifer Inocarpus chestnut) (Tahitian batatas Ipomoea (SweetPotato) 2 sp. Musa (Banana) methysticum Piper (Kava) sp. Pteridium (Bracken) tuberosum Solanum CONTAMINATION (WhitePotato) dulcis Spondias Ambarella) Apple(Otaheiteor sagittifolium Xanthosoma Aroid Taro) Ear Elephant (Arrowleaf CONTAMINATION 40-45 3 1 50-55 3 3 60-65 3 5 70-75 3 7 80-85 4 9 90-95 5 11 100-105 4 13

Table 8.19 Levels of confidence from DFA classification of archaeological starch from Heketa TP3. NB High confidence (black), moderated confidence (medium grey) and low confidence (light grey)

Depth Layer Spit macrorrhiza Alocasia Aroid Taro) (Giant paeoniifolius Amorphophallus Aroid Yam) Foot (Elephant altilis Artocarpus (Breadfruit) esculenta Colocasia Aroid Taro) (Common merkusii Cyrtosperma Aroid Taro) Swamp (Giant fagifer Inocarpus chestnut) (Tahitian batatas Ipomoea (SweetPotato) 2 sp. Musa (Banana/plantain) methysticum Piper (Kava) dulcis Spondias Ambarella) Apple(Otaheiteor 40-45 3 1 50-55 3 3 60-65 3 5 70-75 3 7 80-85 4 9 90-95 5 11 100-105 4 13

Table 8.20 Final table documenting species represented by archaeological starch within Heketa TP3

From this list of classifications, a final table of species identified in Heketa TP3 was produced. The distribution of these classifications within the three sampled cultural deposits indicates some variation among the types of species present, but very little variation in the number of species. Layer 3 (40-75cmbs) contains all four of the aroids— A. macrorrhiza, A. paeoniifolius, C. esculenta and C. merkusii, P. methysticum, several arboreal species including

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A. altilis, I. fagifer, S. dulcis, and one probable contaminant- I. batatas. Layer 4 (80-90, and 100-105cmbs) contains all of the aroids apart from C.merkusii, a range of arboreal species including A. altilis, I. fagifer, and Musa sp. 2, and also I. batatas. Layer 5 represents an ash lens within Layer 4, and therefore was expected to contain very little, if any starch. During processing only a few grains were extracted from this deposit, and only one confident classification could be assigned to these grains during this final stage of analysis, being I. fagifer.

Figure 8.5 Archaeological and reference starch: (A) archaeological starch identified as Artocarpus altilis, (B) modern starch of A. altilis, (C) archaeological starch identified as Alocasia macrorrhiza, (D) modern starch of A. macrorrhiza, (E) archaeological starch identified as Amorphophallus paeoniifolius, (F) modern starch of A. paeoniifolius, (G) archaeological starch identified as Barringtonia asiatica, (H) modern starch of B. asiatica, (I) archaeological starch identified as Colocasia esculenta, (J) modern starch of C. esculenta, (K) archaeological starch identified as Curcuma longa, (L) modern starch of C. longa, (M) archaeological starch identified as Cyrtosperma merkusii, (N) modern starch of C. merkusii, (O) archaeological starch identified as Dioscorea alata, (P) modern starch of D. alata.

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Figure 8.6 Archaeological and reference starch cont.: (Q) archaeological starch identified as Dioscorea bulbifera (R) modern starch of D. bulbifera, (S) archaeological starch identified as Dioscorea esculenta, (T) modern starch of D. esculenta, (U) archaeological starch identified as Dioscorea nummularia, (V) modern starch of D. nummularia, (W) archaeological starch identified as Inocarpus fagifer, (X) modern starch of I. fagifer, (Y) archaeological starch identified as Ipomoea batatas, (Z) modern starch of I. batatas, (AA) archaeological starch identified as Musa sp., (AB) modern starch of Musa sp., (AC) archaeological starch identified as Piper methysticum, (AD) modern starch of P. methysticum, (AE) archaeological starch (contaminant) identified as Solanum tuberosum, (AF) modern starch of S. tuberosum, (AG) archaeological starch identified as Spondias dulcis, (L) modern starch of S. dulcis.

Comparison of modern Pacific production systems Comparative ethnographic examples from the Western Pacific region were analysed to establish the range of plant species exploited through production or collection techniques. Nutritional, labour, and productivity data were collated from published examples, to assess and compare the nature of decision-making in a variety of production systems. These variables will be compared and contrasted to discuss decision-making and crop selection. Five systems from the Western Pacific were chosen due to the nature of comparable data available from modern ethnographic studies. These included the Gadio Enga of the New Guinea Highlands (Dornstreich 1977), 160

Tongatapu (Ministry of Agriculture and Forestry 2001), Bellona Island (Christiansen 1975), Ontong Java (Bayliss-Smith 1973, 1977, 1986) and Anuta (Yen 1973b) in the Solomon Island Outliers. These systems were chosen based on the quality and consistency (in terms of time frame for data collection) of data available within each ethnographic study, and to represent a variety of environmental contexts (atoll, high island, raised limestone islands). Several systems also involved some use of cash-cropping (Bellona, Ontong Java and Tongatapu), but it will be argued here that these cannot be easily separated from subsistence production as these contribute to total efficiency (total de-facto output divided by primary and secondary inputs) (Bayliss-Smith 1977) within these systems.

Data was collated on the range of plant species cultivated within each system, and the labour inputs and yields for each of these. Units of labour inputs were recorded as hours invested in agricultural production and collection of gathered species, specifically those invested in garden preparation, maintenance and harvesting of crops. These were compared to outputs in terms of yield in kilos and nutritional benefits over the same time frame. These figures were used to calculate basic efficiency or rate of return ratios for each of these species in terms of nutritional and yield outputs to labour investment inputs. Understanding the basic energetic inputs to output ratios of these systems, without consideration of variables such as social values, acreage planted or harvested, seasonal variation and single or multi-cropping techniques is an important first step towards detailing the nutritional costs and benefits each system within an Human Ecological framework. While it is important to acknowledge the role that other variables play in decision-making, it is not possible to create a model that incorporates all of them. These variables cannot be determined archaeologically, and so providing basic quantifiable nutritional, labour and productivity data may at least allow interpretation of the past through modelling hypothetical and testable production systems.

Nutrition The exploited plant species listed within each of the ethnographic examples, many of which feature in the archaeobotanical record from of Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO- Nt-2), were assessed for their calorific, protein, fat and carbohydrate values for 100g of edible plant material. Some spices and ceremonial or medicinal plants such as kava (Piper methysticum) and betel nut (Areca catechu) were also included in this analysis, primarily because these were cultivated and could have contributed to dietary needs, even if that was not their primary purpose. Nutritional data was collected and collated for these species, allowing comparisons to be made across each system. A total nutritional figure per 100g for each species was created that added all of the values together. These species were considered separately, and then as groups described by their authors as horticultural, semi-cultivated, or gathered (or close synonyms of these such as gardening). These nutritional values are heavily relied upon in the literature on Evolutionary Ecology and Human Ecology to predict subsistence choices, and 161 provide interesting comparison with ethnographically recorded cases of labour and energy investment.

Gadio Enga, New Guinea Of the modern systems analysed, the largest range of utilised plant species was present within the Gadio Enga system observed by Dornstreich (1973, 1977) in Highland New Guinea. A total of 35 species was recorded over an entire subsistence season (one year). Animal foods were included in this dataset and contributed to 25% of dietary protein and 75% of fats, but were not included in this modelling exercise which targets the role of plants within diet. A range of nutritional data was collated for species, and the mean of these ranges was used as a final figure for comparison. The species were broken down into groups including horticulture, gathered, sago and silviculture—which involves controlling the establishment, growth, composition, health and quality of forests to meet cultural needs. Interestingly, those species with the highest calorific value for 100g of edible plant were many of the arboreal species that were either managed through silviculture or gathered (see Table 8.21). These included semi-cultivated Pandanus spp., and gathered Canarium spp., Macaranga spp., and Elaeocarpus sp. Root crops grown through horticulture such as Colocasia esculenta, Dioscorea spp. and Ipomoea batatas were ranked 11th, 12th and 15th according to calorific value within this system, where highest calorific value ranked 1 and lowest value ranked 35. In terms of protein value within the same quantity of edible plant, a similar pattern within the Gadio Enga system is evident, although the top ranked species for protein is the introduced Abelmoschus manihot, a horticulturally produced plant. Aside from this cultigen and Phaseolus vulgaris, the horticultural species generally rank very low compared to the silvicultural and gathered arboreal species.

Arboreal species also rank very highly in fat content, comprising the top eight-ranked species within the system. The highest ranked horticultural species is P. vulgaris which was ranked 9th, followed by A. manihot at 10th. The primary root crops are scattered throughout the rankings below these, with Colocasia esculenta ranked 16th, Dioscorea spp. ranked 26th and Ipomoea batatas ranked 28th. Patterning within the ranking of species for carbohydrate value differs from those seen within the other nutritional values. Metroxylon spp. (sago) pith has the highest amount of carbohydrates per 100g within the Gadio Enga system. Several arboreal fruits rank just below this crop, but root crops such as C. esculenta (ranked 4th) rank higher than that seen within the other values.

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Species Mean kcal Rank kcal Protein mean Rank protein Fat mean Rank Fat Carbohydrates mean Rank carbohydrates Total figures Overall rank nutrition Abelmoschus manihot 369.6 5 21.1 1 2.8 10 15.002 14 408.5 5 Areca catechu 249.0 8 5.0 10 4.0 7 47 2 305.0 8 Artocarpus altilis flesh 82.5 16 2.6 16 0.9 11 19.0 12 105.0 17 Artocarpus altilis seeds 150.0 9 6.0 9 0.5 14 30.0 5 186.5 10 Canarium spp. 644.0 2 14.2 5 68.5 1 5.5 27 732.2 2 Carica papaya 43.0 22 0.5 32 0.3 18 11.0 18 54.8 23 Colocasia esculenta 132.5 11 1.7 21 0.4 16 31.6 4 166.2 11 Cucumis sativa 16.0 29 0.6 30 0.1 28 3.6 29 20.3 33 Curcubita moschata 45.0 20 1.0 26 0.1 28 11.7 17 57.8 22 Cympobogon citratus 25.0 27 2.5 17 0.1 28 0.0 33 27.6 31 Dioscorea spp 128.0 12 1.8 20 0.1 26 26.9 7 156.7 13 Diplazium sp. 341.0 6 14.3 4 0.1 26 8.3 21 363.7 7 Edible ferns 50.0 18 0.3 34 3.2 8 6.0 24 59.5 21 Elaeocarpus sp. 375.0 4 15.0 3 30.0 4 16.0 13 436.0 4 Ficus sp. 13.0 32 7.6 8 9.0 5 5.3 28 34.9 28 Gnetum gnemon 28.0 26 5.0 10 0.2 24 11.0 18 44.2 27 Inocarpus fagifer 240.0 9 4.5 12 4.5 6 40.0 3 289.0 9 Ipomoea batatas 95.0 15 1.6 22 0.1 28 20.1 10 116.8 15 Lagenaria siceraria 14.0 30 0.6 30 0.0 33 3.4 30 18.0 34 Luffa sp. 13.0 32 0.7 29 0.3 17 14.34 15 28.3 30 Macaranga spp. 601.0 3 18.9 2 60.4 3 7.4 22 687.7 3 Manihot esculenta 135.0 10 0.9 28 0.0 33 24.0 8 159.9 12 Metroxylon spp. 323.5 7 0.3 34 0.3 18 79.0 1 403.1 6 Musa spp. 113.5 13 1.2 25 0.3 18 27.3 6 142.3 14 Oenanthe javanica 40.0 23 3.6 15 0.3 18 6.0 24 49.9 25 Pandanus spp. 683.0 1 11.9 6 66.0 2 22.0 9 782.9 1 Pangium edule 40.0 23 1.0 26 0.1 28 20.0 11 61.1 19 Phaseolus vulgaris 97.0 14 8.1 7 3.0 9 0.0 33 108.1 16 Piper betel 44.0 21 4.0 14 0.4 15 6.0 24 54.4 24 Rorippa sp. 17.0 28 2.0 19 0.3 23 3.0 31 22.3 32 Rungia sp. 14.0 30 2.4 18 0.3 18 0.1 32 16.8 35 Saccharum edule 38.0 25 4.1 13 0.2 24 5.5 27 47.8 26 Saccharum officinarum 58.0 17 0.5 32 0.0 33 14.0 16 72.5 18 Setaria palmifolia 22.0 27 1.4 24 0.6 13 6.5 23 30.5 29 Zingiber zerumbet 48.0 19 1.5 23 0.9 11 9.5 20 59.9 20

Table 8.21 Nutritional figures and rankings for species within the Gadio Enga system according to calories, protein, fats, carbohydrates and total nutrition figures (data from Dornstreich 1974, 1978) 163

A total nutritional figure was created by combining the values for calories, protein, fat and carbohydrates for each species from 100g of edible material (see Figure 8.7). This enabled an overall comparison of all species within the system. According to this value, Pandanus spp. had the highest nutritional value with a total figure of 782.9/100g, followed by a number of gathered arboreal species. The highest ranked horticulturally produced crop is again A. manihot, with a figure of 408.5/100g, while root crops such as C. esculenta rank 11th with a figure of 166.2/100g, and Dioscorea spp. rank 13th with a figure of 156.7/100g. When the averages of each of the production groups are compared, sago production ranks the highest; however, this is biased by this group containing only this single species. When the remaining groups are compared, those species grown using silviculture appear to have the highest total nutrition figures with an average of 246.4/100g, followed closely by gathered species which have an average of 225.9/100g. Surprisingly, horticulturally produced species have the lowest average— 99.4/100g within the Gadio Enga system.

Figure 8.7 Nutritional comparison of species within the Gadio Enga plant production system (data from Dornstreich 1974, 1978)

Bellona, Solomon Islands The plant production system on Bellona was recorded by Christiansen (1975) between the years 1965-66, describing a total of 29 plant species utilised within this system. Species within this system were categorised by the author into two groups: horticulture and semi-cultivated. The latter group includes plants which require some anthropogenic manipulation for initial growth but are mostly untended after this time apart from harvesting on a regular basis or opportunistically. This semi-cultivated category includes many arboreal species from which fruits are harvested and mostly eaten raw, or through some processing to remove toxins

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(Christiansen 1975:36-7). Gathering or hunting of plant, animals and fish is described by Christiansen as a means of filling the temporal gap in food supplies during different seasons, during which the bulk of available foods are sourced from outside horticultural practices for at least a month or more.

Firstly, in terms of calories, the highest ranked species are again the arboreal semi- cultivated species such as Pandanus spp. and Canarium spp., which is similar to the rankings seen in the Gadio Enga system (see Table 8.22). Here though, the highest ranked horticultural species is Cocos nucifera (coconut), ranked 3rd with a calorific value of 425.5/100g for the mature meat of the fruit. This is followed by the horticulturally produced root crop Pueraria lobate (kudzu vine), the rhizome of which has a calorific value of 382/100g. Kudzu is not often cultivated today, but instead is naturalised on many Pacific Islands. The common root crops such as Colocasia esculenta (12th) and Dioscorea spp.(14-15th, 17-18th, 23rd) appear within the middle of the spread of calorific rankings along with Musa spp. (16th). Many of the arboreal species of which fruits are eaten raw, such as Spondias dulcis and Syzygium spp., rank the lowest of all the utilised species.

Comparison of protein values reveals a similar pattern to calories. Semi-cultivated arboreal species dominate the highest rankings based on protein content, with the highest ranking horticultural species, Artocarpus altilis (seeds) being ranked 6th. Root crops generally rank very low (>15th), apart from Tacca leontopetaloides (Polynesian arrowroot), which ranks 7th with a protein value of 5.1/100g. The fat content of these species again follows a similar pattern to calories and protein, with C. nucifera ranking the highest of the horticultural species with 34-43/100g (ranked 3-4th), followed by T. leontopetaloides ranked at 8th with 2.6/100g. The remaining root crops all have less than 0.4/100g and rank lower than 15th. Similar to the patterning observed within the Gadio Enga system, the carbohydrate values of these species provides a different distribution of rankings compared to the other nutritional figures. Saccharum officinarum (sugarcane) ranks the highest, with a total value of 100/100g— meaning that this species is entirely composed of carbohydrates. Root crops such as Alocasia macrorrhiza, Colocasia esculenta, Tacca leontopetaloides, Ipomoea batatas and Dioscorea spp. also rank very highly according to carbohydrate value, along with some semi-cultivated arboreal species. These figures are not all that surprising, considering that the carbohydrate value of these species is generally argued to be the reason that they were initially domesticated.

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Species Mean kcal Rank kcal Protein mean Rank protein Fat mean Rank Fat Carbohydrates mean Rank carbohydrates Total figures Overall rank nutrition Alocasia macrorrhiza 106.0 18 2.0 17 0.2 22 22 10 130.1 18 Amorphophallus paeoniifolius 135.0 10 2.2 14 0.1 27 18.19 18 155.5 13 Artocarpus altilis flesh 82.5 22 2.6 12 0.9 12 19.0 17 105.0 22 Artocarpus altilis seeds 150.0 9 6.0 6 0.5 13 30.0 5 186.5 10 Burckella sp. 56.0 25 1.1 25 1.3 11 20.1 12 78.5 24 Canarium spp. 644.0 2 14.2 2 68.5 1 5.5 27 732.2 2 Canavalia sp. 372.0 6 34.8 1 1.5 10 51.7 3 460.0 5 Cocos nucifera immature meat 105.0 20 2.4 13 8.0 7 7.0 26 122.4 19 Cocos nucifera mature meat 425.5 3 4.4 9 43.2 3 10.5 23 483.6 3 Cocos nucifera mature milk 328.5 7 3.4 10 34.5 4 5.5 27 371.9 7 Colocasia esculenta 132.5 12 1.7 21 0.4 15 31.6 4 166.2 11 Cucumis melo 34.0 28 0.8 27 0.2 18 8.0 25 43.0 29 Dioscorea alata 119.7 15 2.1 16 0.1 25 19.6 16 141.5 16 Dioscorea bulbifera 106.0 18 1.9 18 0.1 27 25.3 8 133.3 17 Dioscorea esculenta 120.3 14 1.8 19 0.2 21 20.05 14 142.3 14 Dioscorea nummularia 106.7 17 1.7 22 0.1 29 13.3 20 121.7 20 Dioscorea pentaphylla 82.0 23 1.7 20 0.1 25 20.0 15 103.8 23 Ficus sp. 13.0 31 7.6 5 9.0 6 5.3 29 34.9 30 Gnetum sp. 28.0 29 5.0 8 0.2 18 11.0 22 44.2 28 Ipomoea batatas 95.0 21 1.6 23 0.1 23 20.1 12 116.8 21 Mangifera sp. 60.0 24 0.8 27 0.4 14 15.0 19 76.2 25 Manihot esculenta 135.0 10 0.9 26 0.0 30 24.0 9 159.9 12 Morinda citrifolia 38.0 27 2.7 11 1.8 9 12.0 21 54.5 26 Musa spp. 113.5 16 1.2 24 0.3 16 27.3 6 142.3 15 Pandanus spp. 683.0 1 11.9 3 66.0 2 22.0 10 782.9 1 Pueraria lobata 382.0 4 2.1 15 0.1 23 27.1 7 411.3 6 Saccharum officinarium 375.0 5 0.0 31 0.0 30 100.0 1 475.0 4 Spondias dulcis 43.7 26 0.8 27 0.3 16 3.65 31 48.5 27 Syzygium spp. 21.0 30 0.7 30 0.2 18 5.3 29 27.2 31 Tacca leontopetaloides 122.0 13 5.1 7 2.6 8 89.4 2 219.1 9 Terminalia sp. 258.0 8 9.6 4 24.0 5 8.3 24 299.9 8

Table 8.22 Nutritional figures and rankings for species within Bellona Island system according to calories, protein, fats, carbohydrates and total nutrition figures (data from Christiansen 1975) 166

Finally, the total nutritional figures for all 29 plant species within the Bellona Island system were compared (see Figure 8.8). From these, Pandanus spp. again ranks the highest, followed by Canarium spp., another group of semi-cultivated arboreal species. The highest value of the horticulturally produced species was recorded in C. nucifera mature meat with a total nutritional figure of 483.5/100g. A mixture of cultivated grasses (sugarcane), fruits (Canavalia spp. and C. nucifera mature milk), and root crops (Pueraria lobata) all had total figures greater than 300/100g, and thus were ranked higher than the next highest ranked semi- cultivated species— Terminalia spp. When the two groups are compared, the averages of each are essentially very similar. Horticulturally produced species have an average total nutritional figure of 215.8/100g, while semi-cultivated species have an average of 205.9/100g.

Figure 8.8 Nutritional comparison of species within the Bellona Island plant production system, showing exponential trend lines for horticultural and semi-cultivated taxa (data from Christiansen 1975)

Anuta, Solomon Islands An ethnographic, ecological and archaeological survey of Anuta was carried out by Yen and others (1973b) over several field seasons in 1970-71. Details of Anutan subsistence were recorded over a period of 37 days, with the aim of investigating the reputed intensity of agricultural practice on the island (Yen and Gordon 1973). The use of 19 plant species was recorded within this survey. When the nutritional values of these species are compared, some very different patterning can be observed from both the Gadio Enga and Bellona Island systems, primarily due to the reduced number of exploited arboreal species (see Table 8.23). Unlike the other modern systems, the Anutan plant species were not divided by the authors into any groups. However, since labour and productivity data were recorded for only a few of these, this was taken to indicate that these were primary crops, while all others were most likely

167 supplementary species. This assumption may not be correct, and so the interpretation of these categories within the nutritional figures will not carry the same weight as those used in the other systems.

The archaeology and ecology of Anuta and nearby Tikopia have been compared by researchers in the past (Firth 1939; Kirch 1995; Kirch and Yen 1982). Agricultural production on Tikopia was similar to Anuta in a number of ways including major crop dominance, the use of some tree species as well as root crops, and the mulching of taro and manioc with coconut fronds (Yen and Gordon 1973). Some important differences also existed in the past between these two locations. Sweet potato is significantly more important in Tikopia than Anuta, but is produced alongside other root crops using less intensive agricultural techniques within the mountainous areas where dryland terracing is absent (Firth 1939; Yen and Gordon 1973). Tikopian agriculture was instead more intensive in flat corraline areas where rotational practices with crop successions of taro, manioc or sweet potato predominate. These factors were taken into consideration alongside the availability of labour and yield data when deciding to use data from Anuta, rather than Tikopia. It was decided that, as described by Yen (1973), Anuta represented a highly intensive agricultural system, possibly the most intensive within the Pacific and presented a unique case study for this modelling exercise.

Calorific comparison indicates that the highest ranked species are the few arboreal species within this system, including Canarium spp. with a value of 644/100g, followed by mature meat of Cocos nucifera with a value of 425.5/100g. Those species with little other data recorded, being supplementary species such as Curcuma longa (turmeric), Metroxylon salomonense (sago), and Inocarpus fagifer, are often ranked higher than ‘primary crops’ such as Colocasia esculenta, Manihot esculenta (cassava), and Cyrtosperma merkusii (giant swamp taro). A slightly different range of supplementary crops dominate the highest rankings according to protein content. Canarium spp. again takes out the top rank with a value of 14.2/100g, followed by another fruit tree, Barringtonia procera, with a value of 9.7/100g. These arboreal species are then followed in ranking by a number of root crops (Curcuma longa and Tacca leontopetaloides). The highest ranked ‘primary crop’ is Cocos nucifera with a protein value of 4.35/100g within mature meat. All primary root crops have a protein value of less than 2/100g.

Comparison of the fat content of these plant species indicates that both primary and supplementary arboreal species generally have higher values than root crops. Those that differ from this trend include C. longa with a ranking of 3rd, and T. leontopetaloides with a ranking of 9th out of the 21 total species. Carbohydrate values provide a different trend, but both of these root crops again rank very highly. In fact, T. leontopetaloides has the highest carbohydrate content with a value of 89.4/100g. Primary root crops such as Colocasia esculenta, Dioscorea

168 spp. and Manihot esculenta all range from 20-30/100g and rank in the top ten of the 19 species. Again, this trend was expected.

The overall nutrition figures suggest that in general, those species grouped here as supplementary have higher nutritional value than those grouped as primary crops (Figure 8.9), as expected if production costs are lower for certain crops with low-to-medium nutritional value. The averages of these groups confirm this trend, where supplementary species have an average of 310.5/100g, while primary species only have an average of 190.3/100g. Within this pattern it was also observed that arboreal species tend to have higher values than root crops. The highest overall ranked species was Canarium spp. with a value of 732.2/100g, followed by C. nucifera mature meat with 483.5/100g. The highest ranked root crop was C.longa with a value of 437/100g, and the next was T. leontopetaloides with 219.1/100g. The highest ranked primary root crop was C. esculenta, followed closely by M. esculenta.

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Species Mean kcal Rank kcal Protein mean Rank protein Fat mean Rank Fat Carbohydrates mean Rank carbohydrates Total figures Overall rank nutrition Alocasia macrorrhiza 106.0 16 2.0 12 0.2 17 22 11 130.1 16 Areca catechu 249.0 7 5.0 5 4.0 8 47 4 305.0 7 Artocarpus altilis 82.5 19 2.6 10 0.9 11 19.0 15 105.0 19 Barringtonia procera 243.0 8 9.7 2 11.8 4 35.3 6 299.8 8 Burckella obovata 56.0 20 1.1 17 1.3 10 20.1 12 78.5 20 Canarium sp. 644.0 1 14.2 1 68.5 1 5.5 20 732.2 1 Cocos nucifera immature meat 105.0 17 2.4 11 8.0 6 7.0 18 122.4 17 Cocos nucifera mature meat 425.5 2 4.4 7 43.2 2 10.5 17 483.6 2 Cocos nucifera mature milk 328.5 5 3.4 9 34.5 3 5.5 20 371.9 6 Colocasia esculenta 132.5 11 1.7 14 0.4 13 31.6 7 166.2 11 Curcuma longa 354.0 4 8.0 3 10.0 5 65.0 3 437.0 3 Cyrtosperma merkusii 122.0 13 0.5 19 0.2 16 19.91 14 142.6 14 Dioscorea spp. 128.0 12 1.8 13 0.1 18 26.9 9 156.7 13 Inocarpus fagifer 240.0 9 4.5 6 4.5 7 40.0 5 289.0 9 Ipomoea batatas 95.0 18 1.6 15 0.1 19 20.1 12 116.8 18 Manihot esculenta 135.0 10 0.9 18 0.0 20 24.0 10 159.9 12 Metroxylon salomonense 323.5 6 0.3 21 0.3 14 79.0 2 403.1 4 Musa spp. 113.5 15 1.2 16 0.3 14 27.3 8 142.3 15 Piper betel 44.0 21 4.0 8 0.4 12 6.0 19 54.4 21 Saccharum officinarum 375.0 3 0.5 20 0.0 20 14.0 16 389.5 5 Tacca leontopetaloides 122.0 13 5.1 4 2.6 9 89.4 1 219.1 10

Table 8.23 Nutritional figures and rankings for species within Anutan system according to calories, protein, fats, carbohydrates and total nutrition figures (data from Yen 1973b)

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Figure 8.9 Nutritional comparison of species within the Anutan plant production system, showing exponential trend lines for primary and supplementary taxa (data from Yen 1973b)

Tongatapu, Tonga Archipelago In 2001, an agricultural census was conducted by the Tongan Ministry of Agriculture and Forestry. This included a survey of agricultural holdings, productivity and labour investment, from which a list was provided of all plant species utilised on Tongatapu. This list contains 22 plant species, and was again broken down for comparative purposes into two groups, horticultural and semi-cultivated based on ethnographic data from European Contact-era recordings of subsistence strategies (Beaglehole and Beaglehole 1941; Cook in Beaglehole 1969; Cook 1875; Gifford 1929; La Perouse 1799; La Billardiere 1800; Mariner in Martin 1991; Maude 1965; Wilson 1799; Waldegrave 1873). This census was chosen over Maude’s (1965) research based on the geographic scope (all of Tongatapu) and consistency of labour investment with yield data in terms of the time period of data collection (one year).

The distribution of calorific values of these crops follows a similar trend to most of the other modern systems. Arboreal species including Pandanus spp., Canarium spp. and Cocos nucifera have the highest rankings, while the root crops are ranked below these at 7th (Colocasia esculenta), 8th-11th (Dioscorea spp.) and 10th (Xanthosoma sagittifolium) (see Table 8.24). All three varieties of Musa spp. rank just below these with calorific values ranking from 11th to 13th. Comparison of protein content indicates that arboreal species again take out the top 9 rankings, but within these are a number of horticulturally produced crops such as A. altilis with a protein value of 2.55/100g, and C. nucifera. The highest ranked root crop is A. macrorrhiza with a value of 1.95/100g, followed by C. esculenta with a value of 1.7/100g. The lowest ranked

171 species are semi-cultivated fruit crop S. dulcis and horticulturally produced Piper methysticum (kava), which is cultivated as a sedative for medicinal and recreational consumption, and is therefore not consumed to meet daily nutritional requirements.

The fat content of these 20 species included in this nutritional comparison demonstrate a very similar pattern to that observed within both other systems, and also within the previously described values. Many arboreal species are ranked highest, with fat content values greater than 1/100g. The highest ranked root crop is X. sagittifolium (elephant ear taro) with a value of 0.4/100g, followed by C. esculenta. All three Musa spp. have values from 0.3-0.4/100g and so rank closely with these root crops. Ipomoea batatas, on the other hand, is ranked very low with a value of only 0.1/100g. The ranking of carbohydrate values again demonstrate the high values of horticulturally produced crops such as C. esculenta (31.6/100g), Musa spp. (27.3-32/100g), Dioscorea spp. (19-26.85/100g), and X. sagittifolium (23.63/100g).

When these nutritional figures are combined, it would appear that arboreal species again rank much higher than root crops (see Figure 8.10). Once again Pandanus spp. is ranked the highest, followed by Canarium spp. and C. nucifera. The highest ranked root crop is also C. esculenta, followed by one of the recorded Dioscorea sp. This reflects the overall similarities in the species composition of the Tongan system with the others. When the averages of the two horticultural and semi-cultivated groups are compared, it becomes clear that the semi-cultivated species have greater overall nutritional value than the horticultural species. The average for the semi-cultivated group is 337.9/100g, while that for the horticultural species is much lower at 183.8/100g.

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Species Mean kcal Rank kcal Mean protein Rank protein Fat mean Rank Fat Carbohydrates mean Rank carbohydrates Total figures Overall rank nutrition Alocasia macrorrhiza 106.0 15 2.0 10 0.2 17 22 10 130.1 16 Artocarpus altilis 82.5 19 2.6 7 0.9 8 19.0 14 105.0 19 Artocarpus heterophyllus 95.0 17 1.7 13 0.6 9 23.0 9 120.3 17 Canarium sp. 644.0 2 14.2 1 68.5 1 5.5 19 732.2 2 Cocos nucifera immature meat 105.0 16 3.4 5 34.5 4 5.5 19 148.4 12 Cocos nucifera mature meat 425.5 3 4.4 4 43.2 3 10.5 17 483.6 3 Cocos nucifera mature milk 328.5 5 2.4 8 8.0 5 7.0 18 345.9 5 Colocasia esculenta 132.5 7 1.7 13 0.4 13 31.6 3 166.2 7 Dioscorea sp. 1 128.0 8 1.8 11 0.1 18 26.9 6 156.7 8 Dioscorea sp. 2 122.0 11 2.1 9 0.1 21 19.6 13 143.8 13 Disocorea sp. 3 128.0 8 1.8 11 0.1 18 26.9 6 156.7 8 Inocarpus fagifer 240.0 6 4.5 3 4.5 6 40.0 1 289.0 6 Ipomoea batatas 95.0 17 1.6 15 0.1 20 20.1 12 116.8 18 Morinda citrifolia 38.0 21 2.7 6 1.8 7 12.0 16 54.5 20 Musa sp. 1 122.0 11 1.3 17 0.4 11 32.0 2 155.7 10 Musa sp. 2 113.5 13 1.2 18 0.3 14 27.3 4 142.3 14 Musa sp. 3 113.5 13 1.2 18 0.3 14 27.3 4 142.3 14 Pandanus sp. 683.0 1 11.9 2 66.0 2 22.0 10 782.9 1 Piper methysticum 6.0 22 0.2 22 0.5 10 0.2 22 6.9 22 Saccharum officinarum 375.0 4 0.5 21 0.0 22 14.0 15 389.5 4 Spondias dulcis 43.7 20 0.8 20 0.3 14 3.65 21 48.5 21 Xanthosoma sagittifolium 125.0 10 1.5 16 0.4 11 23.6 8 150.5 11

Table 8.24 Nutritional figures and rankings for species within Tongan system according to calories, protein, fats, carbohydrates and total nutrition figures (data from Ministry of Agriculture and Forestry 2001)

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Figure 8.10 Nutritional comparison of species within the Tongan plant production system, showing exponential trend lines for horticultural and semi-cultivated taxa (data from Ministry of Agriculture and Forestry 2001)

Ontong Java Atoll, Solomon Islands The atoll of Ontong Java was surveyed by Bayliss-Smith (1973, 1977, 1986) from 1970-1971 as a component of his doctoral research and part of the South Pacific Smallholder Project. This survey targeted agricultural production and associated ecosystem manipulation over time, with a focus on the limits of island carrying capacity. Through this, a total of 13 utilised plant species were recorded, including a range of tree crops from which fruits were harvested and also a number of traditional and recently introduced root crops. Most of the atoll is divided into specific ecosystems or vegetation units from which particular food products are extracted. Cash- cropping involving copra production and exports also have enabled other subsistence products to be imported back into the island. These plant species can be roughly divided into two groups, primary and supplementary based on Bayliss-Smith’s (1977) description of production techniques and energy investment. For example, Curcuma longa is grown extensively on the island and is cultivated as a form of social production rather than subsistence. This species is still therefore grouped here as primary rather than a supplementary subsistence crop.

A calorific comparison of species within the Ontong Java system ranks Pandanus tectorius highest, with a value of 683/100g (see Table 8.25). Within this system, Pandanus is cultivated within coconut woodland and is considered a primary crop. Following this in the calorific ranking is Cocos nucifera, and then Saccharum officinarum (sugarcane) which are both primary cultigens. Other arboreal species such as Carica papaya (papaya) and Artocarpus altilis

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(breadfruit) rank at the bottom of the system, while Musa sp. also ranks relatively low at 10th. The highest ranked primary root crop is C. longa, followed closely by C. esculenta and X. sagittifolium. In terms of protein, Pandanus also ranks the highest, and is followed by one primary and one supplementary root crop, C. longa and Tacca leontopetaloides. Interestingly, the next highest ranked primary root crop according to protein value is Alocasia macrorrhiza. Patterning of fat content follows very closely that of protein values, with some minor reshuffling within the rankings. The same four species have the highest rankings— a combination of both arboreal and root crops, and supplementary and primary crops. The lowest ranked species is S. officinarum with a fat value of 0/100g, ranked just below Ipomoea batatas which has a fat value of 0.1/100g. The ranking of these species according to carbohydrate values follows a now well-established pattern, dominated by high-ranked primary and supplementary root crops such as T. leontopetaloides (1st), C. longa (2nd), and C. esculenta (3rd).

The overall nutritional figures reflect the consistently high rankings of arboreal species such as P. tectorius and C. nucifera, as well as supplementary root crops including C. longa and T. leontopetaloides (see Figure 8.11). The primary root crops were ranked 7th and below. When the averages of the two groupings are compared, there is only a slight difference between them. Primary crops have a slightly higher average (265.8/100g) than those species categorised as supplementary (222.8/100g).

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Species Mean kcal Rank kcal Protein mean Rank protein Fat mean Rank Fat Carbohydrates mean Rank carbohydrates Total figures Overall rank nutrition Alocasia macrorrhiza 106.0 11 2.0 8 0.2 13 22 6 130.1 12 Artocarpus altilis 82.5 14 2.6 6 0.9 7 19.0 10 105.0 14 Carica papaya 43.0 15 0.5 14 0.3 11 11.0 12 54.8 15 Cocos nucifera immature meat 105.0 12 3.4 5 34.5 3 5.5 15 148.4 9 Cocos nucifera mature meat 425.5 2 4.4 4 43.2 2 10.5 13 483.6 2 Cocos nucifera mature milk 328.5 5 2.4 7 8.0 5 7.0 14 345.9 5 Colocasia esculenta 132.5 6 1.7 9 0.4 9 31.6 3 166.2 7 Curcuma longa 354.0 4 8.0 2 10.0 4 65.0 2 437.0 3 Cyrtosperma merkusii 122.0 8 0.5 13 0.2 12 19.91 9 142.6 10 Ipomoea batatas 95.0 13 1.6 10 0.1 14 20.1 8 116.8 13 Musa sp. 113.5 10 1.2 12 0.3 10 27.3 4 142.3 11 Pandanus tectorius 683.0 1 11.9 1 66.0 1 22.0 6 782.9 1 Saccharum officinarum 375.0 3 0.5 14 0.0 15 14.0 11 389.5 4 Tacca leontopetaloides 122.0 8 5.1 3 2.6 6 89.4 1 219.1 6 Xanthosoma sagittifolium 125.0 7 1.5 11 0.4 8 23.6 5 150.5 8

Table 8.25 Nutritional figures and rankings for species within Ontong Java system according to calories, protein, fats, carbohydrates and total nutrition figures system (data from Bayliss- Smith 1973, 1986)

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Figure 8.11 Nutritional comparison of species within the Ontong Java plant production system, showing exponential trend lines for primary and supplementary taxa (data from Bayliss-Smith 1973, 1986)

Nutritional comparisons of groupings Some basic statistical analyses were carried out to test whether the overall nutritional figures of species in each grouping or category in each modern system were statistically different. These categories were compared using Student’s two-sample t-test, to evaluate the difference in means between the two categories in light of the pooled standard deviations from both categories (Drennan 1996). The null hypothesis is that the difference between means of the two populations is not greater than 0, while the alternative hypothesis argues that the mean difference is greater than 0. A simple mean comparison may suggest that two groups (or populations) are different, but the t-test indicates how significant that difference actually is. Firstly, the pooled standard deviation and pooled standard error for each system were calculated and then a final figure that indicates the difference between these categories according to standard errors. This figure was then used to explain the probability that the two categories are in fact statistically different using the t test distribution table. Within archaeology the generally accepted probabilty and associated confidence level is 95% or over to statistically reject the null-hypothesis.

Within the Gadio Enga system the horticulture and gathered categories were compared nutritionally (see Table 8.26). The overall nutritional figures of these two categories were shown through Student’s two-sample t-test to have a difference of 1.76 standard errors, arguing that the probability of these groups being from the same population is 20%. Therefore, despite a

177 difference in mean of 126.4, these are not significantly different. The comparison of categories within Bellona Island, Tonga, and Ontong Java all proved that these systems were not statistically different based on the number of species being compared and the associated confidence in statistical differences between nutritional values. The only system that was close to having two statistically different categories was that from Anuta. The comparison of means from primary and supplementary species on Anuta proved that these were nutritionally different with 90% confidence by 1.93 standard errors, but not enough to reject the null hypothesis that these are from the same population.

Nutrition (total figures/100g) Example system Pooled standard deviation Pooled standard error Difference Confidence Mean difference Gadio Enga (horticulture vs gathered) 188.7 71.6 1.8 90% 126.5 Bellona Island (horticulture vs semi-cultivated) 201.5 74.8 0.3 <50% 10.0 Tonga 2001 Agricultural Survey (primary vs supplementary crops) 197.6 94.7 1.6 80% 154.1 Ontong Java (primary vs supplementary crops) 203.6 118.9 0.4 <50% 43.0 Anuta Atoll (primary vs supplementary crops) 153.4 62.0 1.9 90% 120.2

Table 8.26 Statistical comparison of species’ groupings in example systems according to overall nutrition figures/100g

Labour investment Looking beyond the nutritional value of species within each of the example systems, the amount of labour invested in the cultivation, maintenance, harvesting and storage provides a significant insight into the nature of decision-making within these systems. Energy invested into food production can be used as a gauge of where subsistence preferences lie in terms of nutritional outputs, or other social or environmental factors. These labour values will be compared with a variety of these nutritional outputs to discuss diet and productivity within plant food production system.

Gadio Enga, New Guinea Labour investment in this system was recorded by Dornstreich (1973) as instances of activity rather than as accumulated hours or energy spent cultivating and processing plant species. Although an ‘instance of activity’ could equate to any number of man-hours, these data enables at least some basic comparison of labour investment and therefore inputs into the Gadio Enga plant production system. When these figures are compared and then ranked, it becomes clear that labour is directed towards some species and plant categories far more than others.

The highest ranked species by a significant margin is Metroxylon spp., or sago, with a total of 148 recorded instances of activity over the observed subsistence season (see Table 8.27, Figure 8.12). Sago production was noted by Dornstreich (1973, 1977) to provide a significant component of the Gadio Enga diet. Many lower ranked horticultural species share similar values for labour investment, due to the fact that recorded instances of activity were divided between 178 particular resource areas at various altitudes and distances from the village. For this reason, Dioscorea spp. is ranked 2nd with 39.5 instances of activity, followed by a number of cultivated crops such as Abelmoschus manihot, Colocasia esculenta, Cucumis sativa (cucumber), Ipomoea batatas and Manihot esculenta (cassava or tapioca). Many gathered arboreal species such as Ficus sp. (fig), Elaeocarpus sp., Artocarpus altilis and the herb Oenanthe javanica (Japanese parsley) are ranked the lowest within this system according to labour. Most surprising in this data is the very low ranking of A. altilis, as breadfruit is usually cultivated in most Pacific production systems for seeds or flesh and therefore time is invested in initial planting, weeding, harvesting and storage. This ranking may reflect overall or situational subsistence decision- making. For example, that year breadfruit may not have grown well due to environmental factors and so labour was invested elsewhere, or that once breadfruit was established trees required little labour investment.

Species Labour inputs- instances of activity Rank inputs Abelmoschus manihot 30.1 3 Areca catechu 20.7 21 Artocarpus altilis flesh 10.3 26 Artocarpus altilis seeds 10.3 26 Colocasia esculenta 30.1 3 Cucumis sativa 30.1 3 Curcubita moschata 30.1 3 Cympobogon citratus 30.1 3 Dioscorea spp 39.5 2 Edible ferns 28.8 18 Elaeocarpus sp. 11.8 23 Ficus sp. 28.8 18 Ipomoea batatas 30.1 3 Manihot esculenta 30.1 3 Metroxylon spp 148.0 1 Musa spp. 30.1 3 Oenanthe javanica 11.8 23 Pandanus spp. 20.7 21 Pangium edule 11.8 23 Phaseolus vulgaris 30.1 3 Piper betel 30.1 3 Rorippa sp. 30.1 3 Rungia sp. 28.8 18 Saccharum edule 30.1 3 Saccharum officinarum 30.1 3 Setaria palmifolia 30.1 3 Zingiber zerumbet 30.1 3

Table 8.27 Labour investment into species within the Gadio Enga system (data from Dornstreich 1974, 1978)

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Figure 8.12 Labour comparison of species within the Gadio Enga system (data from Dornstreich 1974, 1977)

Bellona, Solomon Islands Christiansen (1975) recorded hours of labour over a year, according to the types of gardens and species within them. Therefore, many species within the Bellona Island system share the same figures for labour investment, and so also have the same rankings when all species are compared (see Table 8.28, Figure 8.13). Despite this, some species stood out within the system. One example of this is the case of Ipomoea batatas which ranked the highest in the system, with around 33,000 hours of labour invested in this crop in a year. Either the cultivation of this species is labour intensive to maintain productive yields, or sweet potato is regarded as a nutritionally or socially important cultigen and therefore time is invested accordingly to increase cultivation more than other horticulturally produced species. Such considerations will be discussed later in the chapter when output to input ratios are calculated. Three Dioscorea spp. are ranked below this, with labour investment values of 16,683 hours/year. Two of the aroids, Alocasia macrorrhiza and Colocasia esculenta rank 5th due to a total of 11,872 hours/year being dedicated to the production of these two cultigens. Horticulturally produced or semi-cultivated arboreal species such as Cocos nucifera, Burckella sp. and Canarium spp. generally rank below these root crops. Artocarpus altilis again ranks the lowest according to labour investment, a trend which requires comparison across these systems.

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Species Labour hrs/yr Rank inputs Alocasia macrorrhiza 11872.0 5 Amorphophallus paeoniifolius 854.1 11 Artocarpus altilis flesh 60.0 27 Artocarpus altilis seeds 60.0 27 Burckella sp. 854.1 11 Canarium spp. 854.1 11 Canavalia sp. 854.1 11 Cocos nucifera immature meat 3333.3 8 Cocos nucifera mature meat 3333.3 8 Cocos nucifera mature milk 3333.3 8 Colocasia esculenta 11872.0 5 Cucumis melo 854.1 11 Dioscorea alata 16683.0 2 Dioscorea bulbifera 854.1 11 Dioscorea esculenta 16683.0 2 Dioscorea nummularia 16683.0 2 Dioscorea pentaphylla 854.1 11 Ficus sp. 854.1 11 Gnetum sp. 854.1 11 Ipomoea batatas 33000.0 1 Mangifera sp. 60.0 27 Manihot esculenta 854.1 11 Morinda citrifolia 854.1 11 Musa spp. 9425.0 7 Pandanus spp. 100.0 26 Pueraria lobata 854.1 11 Saccharum officinarium 854.1 11 Spondias dulcis 60.0 27 Syzygium spp. 60.0 27 Tacca leontopetaloides 854.1 11 Terminalia sp. 854.1 11

Table 8.28 Labour investment into species within the Bellona Is system (data from Christiansen 1975)

Figure 8.13 Labour comparison of species within the Bellona Island system (data from Christiansen 1975)

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Anuta, Solomon Islands Labour investment on Anuta was recorded by Yen (1973b) over a 37 day period for a family group. These figures are compared here according to both species and also grouping (Table 8.29 and Figure 8.14). The highest intensity of labour investment was directed towards horticultural species, with very little investment in supplementary species. This is an expected trend according to the nature and definition of the groupings used within this example system. The highest ranked species according to labour investment is Manihot esculenta with a total of 125.6 recorded hours, followed closely by Colocasia esculenta with 108.6 hours. These two crops have the highest labour investment by a significant margin, with the next ranked species, Artocarpus altilis, having only 62.6 hours dedicated to production. The highest ranked supplementary species are Areca catechu (betel palm) and Piper betel (betel leaf), which are cultivated together for betelnut consumption, each had only 2.2 hours of labour invested over the recorded period. The lower ranked species are all those grouped as supplementary aside from Ipomoea batatas which only had 0.7 hours spent on cultivation and processing.

Species Labour inputs (hrs/37 days) Rank inputs Alocasia macrorrhiza 0.7 12 Areca catechu 2.2 10 Artocarpus altilis 62.6 3 Barringtonia procera 0.7 12 Burckella obovata 24.3 4 Canarium sp. 0.7 12 Cocos nucifera immature meat 10.1 7 Cocos nucifera mature meat 10.1 7 Cocos nucifera mature milk 10.1 7 Colocasia esculenta 108.6 2 Curcuma longa 0.7 12 Cyrtosperma merkusii 18.3 6 Dioscorea spp. 0.7 12 Inocarpus fagifer 0.7 12 Ipomoea batatas 0.7 12 Manihot esculenta 125.6 1 Metroxylon salomonense 0.7 12 Musa spp. 20.3 5 Piper betel 2.2 10 Saccharum officinarum 0.7 12 Tacca leontopetaloides 0.7 12

Table 8.29 Labour investment into species within the Anutan system (data from Yen 1973b)

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Anuta Labour Comparison 140

120

100

Labour inputs Labour (hrs/37 day period) Primary 20 Supplementary

Musa spp. Musa

Piperbetel

Canarium Canarium sp.

Arecacatechu

Dioscoreaspp.

Curcuma longa Curcuma

Ipomoea batatas Ipomoea

Artocarpusaltilis

Inocarpusfagifer

Burckellaobovata

Manihot esculentaManihot

Colocasia esculenta

Barringtonia procera

Alocasia macrorrhiza

Cyrtosperma merkusii

Saccharum officinarum

Tacca leontopetaloides

Metroxylonsalomonense

Cocos nuciferaCocosmature milk

Cocos nuciferaCocosmature meat Cocos nuciferaCocos immature meat Species

Figure 8.14 Labour comparison of species within the Anutan system (data from Yen 1973b)

Tongatapu, Tonga Archipelago During the 2001 agricultural census conducted in Tonga, some data was collected on the amount of labour invested into agricultural production by gender and relationship to the land (owner, family or worker) per week. These labour figures were used to estimate yearly labour investment into agriculture. These total figures were further broken down into estimated species-specific figures through the use of acreage data detailing land dedicated to specific crop production within the same census (see Table 8.30 and Figure 8.15). This avoided dealing with the amount of labour dedicated to most European export crops. The patterning in these figures closely matches that expected based on early historic and ethnographic recordings of Tongan subsistence production. Most labour in the census was invested in the production of the Early yam (Dioscorea sp.2, presumably Dioscorea alata) which is known to be a very important cultigen within Tongan culture and related to surplus production for tributes and festivals. Around 1,122,947.3 hours of labour were directed towards the production of this crop in the census year. Ranked just below this species is another horticulturally produced cultigen, Xanthosoma sagittifolium, which is also a historic introduction into Tonga that is now favoured over the common taro because large quantities of this species can be grown using established dryland production techniques. Due to the popularity of this introduction the common taro ranks 3rd, with 1,015,599.1 hours/year of labour investment. Other root crops also rank highly, including Piper methysticum which ranks 7th with a total of 131,428.5 hours/year of labour investment. Most semi-cultivated species rank below the horticultural crops, aside from 183

Pandanus sp., which ranks above one Dioscorea sp., Saccharum officinarum and A. altilis, with a total of 10,706.9 hours/year investment. It is interesting that Artocarpus altilis once again ranks low within this system.

Species Labour (hrs/yr) Rank inputs Alocasia macrorrhiza 296824.2 6 Artocarpus altilis 5991.6 16 Cocos nucifera immature meat 120503.1 8 Cocos nucifera mature meat 120503.1 8 Cocos nucifera mature milk 120503.1 8 Colocasia esculenta 1015599.1 3 Dioscorea sp. 1 10450.4 14 Dioscorea sp. 2 1122947.3 1 Disocorea sp. 3 461086.1 4 Inocarpus fagifer 3002.1 17 Ipomoea batatas 332514.0 5 Morinda citrifolia 2942.0 18 Musa sp. 1 73029.6 11 Musa sp. 2 56565.4 12 Pandanus sp. 10707.0 13 Piper methysticum 131428.5 7 Saccharum officinarum 7723.8 15 Spondias dulcis 259.7 19 Xanthosoma sagittifolium 1069745.0 2

Table 8.30 Labour investment into species within the Tongan system (data from Ministry of Agriculture and Forestry 2001)

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Figure 8.15 Labour comparison of species within the Tongan system (data from Ministry of Agriculture and Forestry 2001)

Ontong Java, Solomon Islands Energy use and investment of labour within the Ontong Java plant production system over a year was directed primarily towards the cash-cropping and export of copra from Cocos nucifera. This species therefore dominates the man-hours dedicated to agriculture on the island, with a total of 202,960 hours spent cultivating, harvesting and processing this export item over the year recorded by Bayliss-Smith (1973) (see Table 8.31 and Figure 8.16). Within the ethnographic study from which this data was extracted, it is argued that the energy investment into export items rather than subsistence species directly should not be discounted completely as time wasted. The currency generated from these exports enabled other subsistence items to be brought into the Ontong Java system that might not be grown or accessed any other way, and thus still contribute to diet and system outputs, if not directly through a nutritional contribution.

Subsistence items that were grouped as primary crops, such as Colocasia esculenta and Cyrtosperma merkusii, are ranked just below C. nucifera, with around 43,830 hours/year dedicated to each of these species. There is an important tradition of turmeric (Curcuma longa) cultivation on Ontong Java for cultural as well as subsistence reasons, and therefore 9,200 hours/year was spent on the production of this species. Labour investment in all other species was simply described as dedicated toward village crop cultivation or gathering wild fruits, with very low time investment in comparison with the aforementioned major horticultural cultigens.

Species Labour inputs (hrs/yr) Rank inputs Alocasia macrorrhiza 183 7 Artocarpus altilis 183 7 Carica papaya 183 7 Cocos nucifera immature meat 202960 1 Cocos nucifera mature meat 202960 1 Cocos nucifera mature milk 202960 1 Colocasia esculenta 43830 4 Curcuma longa 9200 6 Cyrtosperma merkusii 43830 4 Ipomoea batatas 183 7 Musa sp. 183 7 Pandanus tectorius 183 7 Saccharum officinarum 183 7 Tacca leontopetaloides 183 7 Xanthosoma sagittifolium 183 7

Table 8.31 Labour investment into species within the Ontong Java plant production system (data from Bayliss- Smith 1973, 1986)

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Figure 8.16 Labour comparison of species within the Ontong Java plant production system (data from Bayliss- Smith 1973, 1986)

Labour input comparison of groupings Student’s t-test and associated confidence distribution tables were again used to calculate whether the groupings of species for each modern system used in this study were significantly different according to mean (see Table 8.32). Compared to the levels of confidence in differences generated for the nutritional data, those established for the labour inputs in each system were generally much higher. The groupings or categories within three systems were shown to be statistically different with acceptable confidence. The comparison of labour investment into horticultural and gathered species in the Gadio Enga plant production system was shown to be different by 5.8 standard errors with 99.9% confidence. From this, the absolute means of these groups were shown to be different by a value of 19 hours. Similarly, a comparison of horticultural and semi-cultivated species within the Bellona Island system calculated that these two groupings were different by 2.6 standard errors, the probability of these being different is 98% according to the number of species in each category. Finally, the difference between primary and supplementary species within the Anutan system was shown to be 4.6 standard errors, and so can be argued to be different with 100% confidence. The means of these two categories were different by 14.5 hours. The remaining two systems could not be shown to be statistically different with high probability in labour investment.

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Labour Inputs (hours or instances of activity) Example system Pooled standard deviation Pooled standard error Difference Confidence Mean difference Gadio Enga (horticulture vs gathered) 8.5 3.3 5.8 99.90% 19.0 Bellona Island (horticulture vs semi-cultivated) 649.6 243.1 2.6 98% 619.0 Tonga 2001 Agricultural Survey (primary vs supplementary crops) 6435.6 3086.4 1.9 90% 5889.8 Ontong Java (primary vs supplementary crops) 76265.7 44470.2 1.4 80% 63238.6 Anuta Atoll (primary vs supplementary crops) 10.5 4.6 3.2 100% 14.5

Table 8.32 Statistical comparison of species’ groupings in example systems according to labour input figures

Outputs The nutritional and energy outputs from each of the example systems provide essential data about the costs and benefits of labour investment into each plant species within production systems. These figures were calculated based on the productivity or yield for each species within the time period for which ethnographic data was collected. Because these time periods vary, these figures cannot be compared directly across the different systems. Here, yield and associated outputs will be outlined for each species within each Pacific production system. Because of biases in the data for recorded outputs, the groupings in each example system will not be statistically compared. The section following this discussion on system outputs will focus on patterning in the output to input ratios for each species in each of the example systems. These ratios can be used as a gauge for ‘system efficiency’ according to energy returns.

Gadio Enga, New Guinea System output data for the Gadio Enga was recorded over one subsistence season, or a full year. When the yields for species within this system are directly compared, it becomes clear that the horticulturally produced species have the greatest productivity (see Table 8.33, Figure 8.17). The highest yielding species within this season was Colocasia esculenta, from which 1987.4 kg of edible material was harvested. Ranked just below this was another staple crop, Metroxylon spp., with a total yield of 1117.4kg. Other arboreal species generally ranked low within the system, aside from Musa spp. (although these are technically herbs not trees) which rank 4th as this genus is more like root crops in terms of growth and yields. The highest supplementary arboreal species was Pandanus spp. which was ranked 13th with a total yield of 28.5kg. Root crops such as C. esculenta, Ipomoea batatas, and Manihot esculenta, on the other hand, generally rank relatively high. Dioscorea spp. are the only taxa to differ from this trend, with a ranking of 14th due to a yield of 13.3kg. It is interesting to note the high yield and associated rank of edible ferns, which comes in at 7th with a yield of 60.7kg. From these data, the total yields from each of the groupings revealed an expected pattern. Horticultural production had a total yield of 2935.1kg, while sago had 1117.4kg, gathering had 220.88kg, and finally silviculture had only 50kg.

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Species Yield (kg) Rank Kcal Protein Fats Carbohydrates Abelmoschus manihot 50.0 9 184.8 10558.8 1395.7 7499.4 Areca catechu 0.7 24 1.8 36.2 29.0 340.7 Artocarpus altilis flesh 10.3 17 8.5 263.9 93.1 1966.3 Artocarpus altilis seeds 10.3 17 15.5 620.9 51.7 3104.6 Colocasia esculenta 1987.4 1 2633.4 33786.5 6956.0 628030.7 Cucumis sativa 4.1 21 0.7 24.7 4.1 148.2 Curcubita moschata 84.0 5 37.8 839.9 84.0 9818.0 Cympobogon citratus 0.4 25 0.1 9.1 0.4 0.0 Dioscorea spp 13.4 14 17.1 234.0 16.7 3589.9 Edible ferns 60.7 7 30.3 182.1 1942.3 3641.8 Elaeocarpus sp. 12.1 16 45.5 1821.9 3643.8 1943.4 Ficus sp. 7.0 19 0.9 532.4 628.5 371.8 Ipomoea batatas 498.2 3 473.3 7971.4 498.2 100140.1 Manihot esculenta 62.5 6 84.4 562.6 0.0 15003.4 Metroxylon spp 1117.4 2 3614.9 3352.3 3352.3 882217.1 Musa spp. 172.3 4 195.6 1981.9 517.0 47049.3 Oenanthe javanica 12.5 15 5.0 449.3 37.4 748.8 Pandanus spp. 28.5 13 194.8 3394.2 18825.1 6275.0 Pangium edule 59.0 8 23.6 590.1 59.0 11802.3 Phaseolus vulgaris 0.2 26 0.2 17.4 6.5 0.0 Piper betel 3.1 22 1.4 124.9 12.5 187.4 Rorippa sp. 44.2 11 7.5 883.4 110.4 1325.2 Rungia sp. 7.0 19 1.0 168.3 21.0 3.5 Saccharum edule 0.1 27 0.0 2.3 0.1 3.1 Saccharum officinarum 29.2 12 16.9 145.9 0.0 4085.8 Setaria palmifolia 47.7 10 10.5 668.1 286.3 3102.0 Zingiber zerumbet 0.8 23 0.4 12.1 7.3 76.8

Table 8.33 Output comparison of species in Gadio Enga plant production system (data from Dornstreich 1974, 1978)

Figure 8.17 Output comparison according to yield for species within Gadio Enga system (data from Dornstreich 1974, 1978)

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Bellona, Solomon Islands The yield (mT) for each species within the Bellona system was recorded over a single year. The highest ranked amongst these were three of the yam species— D. alata, D. esculenta and D. nummularia— each with yields of around 211.7mT over the recorded year (see Table 8.34, Figure 8.18). Musa spp. ranked 4th with a yield just below these species. This genus represented the highest ranked arboreal taxa within the system, followed by Cocos nucifera with a yield of 100mT. A number of horticultural root crops such as Ipomoea batatas. Colocasia esculenta, and Alocasia macrorrhiza rank 8-9th with yields from 65-98mT. Some semi-cultivated arboreal species exploited for fruits, including Mangifera sp., Spondias dulcis and Syzygium sp., are ranked 11th with yields of 5mT. The remaining taxa within the system had low yields, and included a range of both horticultural and semi-cultivated species. Overall, the horticultural species collectively had a much higher yield than the range of semi-cultivated species, with a total of 1387.6mT compared to 37.5mT.

Species Yield (mT) Rank Kcal Protein Fat Carbohydrates Alocasia macrorrhiza 65 9 68.9 1267.5 97.5 14300.0 Amorphophallus paeoniifolius 2.5 14 3.4 56.0 1.5 454.8 Artocarpus altilis flesh 2.5 14 2.1 63.8 22.5 475.0 Artocarpus altilis seeds 2.5 14 3.8 150.0 12.5 750.0 Burckella sp. 2.5 14 1.4 27.5 32.5 502.5 Canarium spp. 2.5 14 16.1 355.0 1712.5 137.5 Canavalia sp. 2.5 14 9.3 870.0 37.5 1291.3 Cocos nucifera immature meat 100 5 105.0 2350.0 8000.0 7000.0 Cocos nucifera mature meat 100 5 425.5 4350.0 43200.0 10500.0 Cocos nucifera mature milk 100 5 328.5 3400.0 34500.0 5500.0 Colocasia esculenta 65 9 86.1 1105.0 227.5 20540.0 Cucumis melo 2.5 14 0.9 20.0 5.0 200.0 Dioscorea alata 211.7 1 253.4 4361.0 169.4 41514.4 Dioscorea bulbifera 2.5 14 2.7 48.5 1.5 631.5 Dioscorea esculenta 211.7 1 254.7 3747.1 381.1 42445.9 Dioscorea nummularia 211.7 1 225.9 3493.1 105.9 28198.4 Dioscorea pentaphylla 2.5 14 2.1 43.3 2.0 500.0 Ficus sp. 2.5 14 0.3 189.8 224.0 132.5 Gnetum sp. 2.5 14 0.7 125.0 5.0 275.0 Ipomoea batatas 98 8 93.1 1568.0 98.0 19698.0 Mangifera sp. 5 11 3.0 40.0 20.0 750.0 Manihot esculenta 2.5 14 3.4 22.5 0.0 600.0 Morinda citrifolia 2.5 14 1.0 67.5 45.0 300.0 Musa spp. 202 4 229.3 2323.0 606.0 55146.0 Pandanus spp. 2.5 14 17.1 297.5 1650.0 550.0 Pueraria lobata 2.5 14 9.6 52.5 2.5 677.5 Saccharum officinarium 2.5 14 9.4 0.0 0.0 2500.0 Spondias dulcis 5 11 2.2 40.0 15.0 182.5 Syzygium spp. 5 11 1.1 35.0 10.0 265.0 Tacca leontopetaloides 2.5 14 3.1 127.5 64.5 2235.0 Terminalia sp. 2.5 14 6.5 240.0 600.0 207.5

Table 8.34 Output comparison of species in Bellona Island system (data from Christiansen 1975)

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Figure 8.18 Output comparison according to yield for species within Bellona system (data from Christiansen 1975)

Anuta, Solomon Islands The outputs of the family garden within the Anutan production system, as recorded by Yen (1973b) over a 37 day period, were solely derived from primary species. Clearly there is a bias within the nature of recorded outputs in terms of yield over this time period. Perhaps the season within which these outputs were recorded was that when only a small number of species were being harvested. Another option is that the recorders were only given access to data from particular species. No explanation of the data is given by the authors. In any case, outputs in terms of yield were recorded for only six of the total 19 species. Highest ranked amongst these species is Manihot esculenta, with a total yield of 254.7kg over the recorded period (see Table 8.35, Figure 8.19). There is a significant difference between this crop and the yield of the next highest ranked species, Colocasia esculenta, which had 163.3kg. The remaining taxa all had yields of less than 50kg. Based on these data, the recorders were able to make estimates for the expected annual yield of these species from the number of plants or trees within the plantation. Manioc or Manihot esculenta was expected to have the highest yield of around 2,511kg, followed by C. esculenta with 1610kg. Breadfruit (Artocarpus altilis) was expected to yield around 326kg, while Musa spp., was expected to yield only 127kg over a year. Lowest expected annual yield was from Burckella sp. with only 41kg.

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Species Yield (kg) Rank Kcal Protein Fat Carbohydrates Alocasia macrorrhiza 0.0 0 0.0 0.0 0.0 0.0 Areca catechu 0.0 0 0.0 0.0 0.0 0.0 Artocarpus altilis 33.1 3 27276.9 843.1 297.6 6281.9 Barringtonia procera 0.0 0 0.0 0.0 0.0 0.0 Burckella obovata 4.2 6 2328.6 45.7 54.1 835.8 Canarium sp. 0.0 0 0.0 0.0 0.0 0.0 Cocos nucifera immature meat 0.0 0 0.0 0.0 0.0 0.0 Cocos nucifera mature meat 0.0 0 0.0 0.0 0.0 0.0 Cocos nucifera mature milk 0.0 0 0.0 0.0 0.0 0.0 Colocasia esculenta 163.3 2 216354.0 2775.9 571.5 51598.4 Curcuma longa 0.0 0 0.0 0.0 0.0 0.0 Cyrtosperma merkusii 20.5 4 24993.9 104.5 32.8 4078.9 Dioscorea spp. 0.0 0 0.0 0.0 0.0 0.0 Inocarpus fagifer 0.0 0 0.0 0.0 0.0 0.0 Ipomoea batatas 0.0 0 0.0 0.0 0.0 0.0 Manihot esculenta 254.7 1 343798.2 2292.0 0.0 61119.7 Metroxylon salomonense 0.0 0 0.0 0.0 0.0 0.0 Musa spp. 12.9 5 14619.2 148.1 38.6 3516.3 Piper betel 0.0 0 0.0 0.0 0.0 0.0 Saccharum officinarum 0.0 0 0.0 0.0 0.0 0.0 Tacca leontopetaloides 0.0 0 0.0 0.0 0.0 0.0

Table 8.35 Output comparison of species in Anutan system (data from Yen 1973b)

Anuta Output Comparison 300

250

200

150

100 Outputs (yield (kg)) Primary crops 50 Supplementary

Musa spp. Musa

Artocarpusaltilis

Burckellaobovata

Manihot Manihot esculenta

Colocasia esculenta Cyrtosperma merkusii Species

Figure 8.19 Output comparison according to yield for species within the Anutan system (data from Yen 1973b)

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Tongatapu, Tongan Archipelago The output data within the 2001 Agricultural Census in Tonga had similar biases to that observed of the Anutan system. Data from Tongatapu only included yields of the core cultigens, which have been grouped here as horticultural species. There is therefore a recording bias towards this category. Aside from these core cultigens, many other horticultural and all supplementary species were excluded from the ranking based on outputs. It is not likely that these species had no yield over this time period on Tongatapu, and so to rank these species as ‘0’ for yield would skew the comparisons.

Of those taxa for which there are data, Xanthosoma sagittifolium is ranked the highest with an annual yield of 13,677.5mT on Tongatapu (see Table 8.36, Figure 8.20). This is not unexpected due to the popularity in Tonga of this introduced dryland taro from South America. Ranked just below this with a yield of 13,637.9mT, is the ‘Early yam’ or Dioscorea sp. 2. Again the high yield of this species is not unexpected due to the cultural significance of this cultigen in Tonga. The common taro, Colocasia esculenta, had a recorded annual yield of 9,088.7mT, and was ranked 3rd. Root crops dominate the remaining rankings according to yield, although some arboreal species are interspersed amongst these, including Musa spp. ranked 7-8th and Pandanus sp. ranked 9th with a total annual yield of 1,394.5mT.

Species Yield (mt) Rank Kcal Protein Fat Carbohydrates Alocasia macrorrhiza 1869.0 6 1981.1 36445.0 2803.5 411174.8 Artocarpus altilis 0.0 0 0.0 0.0 0.0 0.0 Artocarpus heterophyllus 0.0 0 0.0 0.0 0.0 0.0 Canarium sp. 0.0 0 0.0 0.0 0.0 0.0 Cocos nucifera immature meat 0.0 0 0.0 0.0 0.0 0.0 Cocos nucifera mature meat 0.0 0 0.0 0.0 0.0 0.0 Cocos nucifera mature milk 0.0 0 0.0 0.0 0.0 0.0 Colocasia esculenta 9088.7 3 12042.5 154507.8 31810.4 2872027.5 Dioscorea sp. 1 126.9 10 162.5 2221.1 158.6 34077.5 Dioscorea sp. 2 13637.9 2 16638.3 280941.1 10910.3 2674395.3 Disocorea sp. 3 5599.8 4 7167.7 97996.1 6999.7 1503539.9 Inocarpus fagifer 0.0 0 0.0 0.0 0.0 0.0 Ipomoea batatas 5100.9 5 4845.9 81614.4 5100.9 1025280.9 Morinda citrifolia 0.0 0 0.0 0.0 0.0 0.0 Musa sp. 1 1833.8 7 2237.2 23839.5 7335.2 586819.4 Musa sp. 2 1632.2 8 1852.6 18770.6 4896.7 445598.7 Musa sp. 3 0.0 0 0.0 0.0 0.0 0.0 Pandanus sp. 1394.5 9 9524.2 165941.6 920348.4 306782.8 Piper methysticum 0.0 0 0.0 0.0 0.0 0.0 Saccharum officinarum 0.0 0 0.0 0.0 0.0 0.0 Spondias dulcis 0.0 0 0.0 0.0 0.0 0.0 Xanthosoma sagittifolium 13677.5 1 17096.9 199691.8 54710.1 3231998.4

Table 8.36 Output comparison of species in Tongan system (data from Ministry of Agriculture and Forestry 2001)

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Figure 8.20 Output comparison according to yield for species within the Tongan system (data from Ministry of Agriculture and Forestry 2001)

Ontong Java, Solomon Islands Data recording caused some issues with the comparison of outputs from species within the Ontong Java plant production system. Four species had no yield data recorded, including all three of the species classified as supplementary. This biases the grouping comparison towards the primary species, but it is not likely that this reflects the actual yield data for the recorded time period. One reason is because of the social importance of turmeric (Curcuma longa) production mentioned by Bayliss-Smith (1973). Despite this, no yield data are given. Perhaps environmental limitations affected production that year.

For those species for which yield data is available, Cocos nucifera, ranks the highest with a total annual yield of 110,700kg (see Table 8.37, Figure 8.21). Of the purely subsistence crops, Cyrtosperma merkusii ranks the highest with an annual yield of 69,230kg. This is followed by another aroid, Colocasia esculenta, with a yield of 40,180kg. These two species are described as core cultigens within the Ontong Java system, grown using wetland techniques that make the most of the island’s geographic and environmental constraints. Several arboreal species such as Carica papaya, Musa spp. and Pandanus tectorius all rank above the lowest ranked root crop, Xanthosoma sagittifolium, which had an annual yield of only 250kg. Perhaps this dryland species of taro is not as viable as wetland varieties within the atoll environment.

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Species Yield (mT) Rank Kcal Protein Fat Carbohydrates Alocasia macrorrhiza 0.0 0 0.0 0.0 0.0 0.0 Artocarpus altilis 0.0 0 0.0 0.0 0.0 0.0 Carica papaya 9.6 6 4.1 47.9 28.7 1052.7 Cocos nucifera immature meat 110.7 1 143.7 4654.6 47230.5 7529.5 Cocos nucifera mature meat 110.7 1 157.0 1605.2 15940.8 3874.5 Cocos nucifera mature milk 110.7 1 121.2 867.2 2952.0 2583.0 Colocasia esculenta 40.2 5 53.2 683.1 140.6 12696.9 Curcuma longa 0.0 0 0.0 0.0 0.0 0.0 Cyrtosperma merkusii 69.2 4 84.5 353.1 110.8 13783.7 Ipomoea batatas 4.7 7 4.5 75.2 4.7 944.7 Musa sp. 1.8 8 2.0 20.2 5.3 480.5 Pandanus tectorius 1.3 9 8.9 155.9 864.6 288.2 Saccharum officinarum 0.4 10 1.5 2.1 0.0 57.4 Tacca leontopetaloides 0.0 0 0.0 0.0 0.0 0.0 Xanthosoma sagittifolium 0.3 11 0.3 3.7 1.0 59.1

Table 8.37 Output comparison of species in Ontong Java production system (data from Bayliss-Smith 1973, 1986)

Figure 8.21 Output comparison according to yield for species within the Ontong Java production system (data from Bayliss-Smith 1973, 1986)

Output to input ratios: Efficiency calculation Once the figures for nutritional values, labour inputs, outputs in terms of yield and associated nutritional returns have been calculated for each species within each system, the ‘efficiency’ of energy returns can then be calculated. In this study, it was deemed appropriate to focus on the species that have been identified archaeobotanically at Talasiu (TO-Mu-2), Leka (J17) and the Heketa (TO-Nt-2) through extracted starch grains and charred macrobotanical remains. This

194 decision was made firstly due to the large range of species within each system, and secondly as the ultimate aim of this analysis is to assess the variation of outputs to inputs within a range of production systems, in order to model archaeological systems. Each of the archaeological species was therefore compared across each of the example systems according to energy and nutritional outputs per time unit. Firstly, a ratio is created for each species that compares the outputs to inputs in terms of yield in kilos, calories, protein, fats, and carbohydrates separately. These are then combined to assess efficiency in terms of the nutritional value of yield per time unit of labour invested. Efficiency is calculated here as:

푌𝑖푒푙푑 (푘푔) × 푛푢푡푟𝑖푡𝑖표푛푎푙 푣푎푙푢푒 (푐푎푙표푟𝑖푒푠, 푝푟표푡푒𝑖푛, 푓푎푡푠 표푟 푐푎푟푏표푦ℎ푦푑푟푎푡푒푠/푘푔)

퐿푎푏표푢푟 (ℎ푟푠 표푟 𝑖푛푠푡푎푛푐푒 표푓 푎푐푡𝑖푣𝑖푡푦)

It is important to point out that the scale of production across each of the example systems is different, but the creation of a simple ratio is argued here to be one technique that diminishes this problem. A larger scale system such as that recorded for Tongatapu requires more hours of labour to gain more outputs, while a smaller scale system such as that on Anuta, which is that of a single family, requires fewer hours for fewer outputs. The calculated ratio is assumed to be the same if production techniques and environmental limitations are the same. One factor that does have to be considered is that the time unit for the Gadio Enga system was recorded as ‘instances of activity’ rather than hours, and so the ratios are going to be skewed towards an over-representation of outputs to inputs. Unfortunately, this cannot be avoided within these comparisons. The results of ratio comparisons within each system will be noted here, and overall patterning between systems discussed within the following section. The ratios for species in each system is compared to gauge the source of highest energy and nutritional returns from labour investment in terms of calories, proteins, fats and carbohydrates. These will also be considered in comparison to the direct yield ratios.

Yield ratio comparison across systems The yield for each species varied dramatically across the different modern systems (see Table 8.38). Some systems such as Anuta and Ontong Java were consistently low-yielding in relation to time invested, thus creating greater disparity across the systems for all species apart from Cyrtosperma merkusii and Ipomoea batatas. The overall greatest difference in yield was that observed for Colocasia esculenta. The highest yielding system for this species was the Gadio Enga, producing an output ratio of 66.1kg per instance of activity, while the lowest yielding system, Ontong Java, produced only 900g per hour. Similarly, Artocarpus altilis yielded 41.7kg/hr within the Bellona system, but only 1kg/instance of activity within the Gadio Enga and 500g/hr within the Anutan production system. Likewise, in assessing the yield ratios for Cocos nucifera, there is a big contrast between the high ratio observed within the Bellona system (30kg/hr) and the low ratio within the Ontong Java example. Many species only had one

195 yield ratio due to the lack of data from other systems. These include: Amorphophallus paeoniifolius, Spondias dulcis and Zingiber zerumbet.

The Bellonan and Tongan systems often had similar high yield ratios as both are raised limestone islands. All of the main Dioscorea yams were relatively high yielding in comparison to labour inputs in the Bellona and Tongan production systems, ranging from 12.14-12.69kg/hr, but had very low ratios in the Gadio Enga system. Similarly, both Musa sp. had yield ratios of 21-25kg/hr, compared to 9.6kg/h in Ontong Java, 5.7kg/instance of activity in Gadio Enga, and only 0.6kg/hr in the Anutan system. Alocasia macrorrhiza was only present in the Bellona and Tongan systems, and had similar yield ratios, 5.48kg/hr in the Bellona system and 6.3kg/hr in the Tongan system. One of the only species for which these systems had radically different ratios, was Ipomoea batatas. The Tongan system produced a yield ratio of 15.3kg/hr, while the Bellona system only produced 3kg/hr. Similarly, Dioscorea bulbifera, commonly a naturalised yam species, was produced in the Tongan system at 12.1kg/hr, while in the Bellona system only 3kg/hr was produced. It should be noted that the ratio within the Tongan system was calculated based on the same yields for all yam varieties, while the Bellona system has specific figures for each Dioscorea species.

Species Gadio Enga Bellona Is Anuta Tongatapu Ontong Java Alocasia macrorrhiza 0.0 5.5 0.0 6.3 0.0 Amorphophallus paeoniifolius 0.0 2.9 0.0 0.0 0.0 Artocarpus altilis 1.0 41.7 0.5 0.0 0.0 Cocos nucifera 0.0 30.0 0.0 0.0 0.5 Colocasia esculenta 66.1 5.5 1.5 8.9 0.9 Curcuma longa 0.0 0.0 0.0 0.0 0.0 Cyrtosperma merkusii 0.0 0.0 1.1 0.0 1.6 Dioscorea alata 0.3 12.7 0.0 12.1 0.0 Dioscorea bulbifera 0.3 2.9 0.0 12.1 0.0 Dioscorea esculenta 0.3 12.7 0.0 12.1 0.0 Dioscorea nummularia 0.3 12.7 0.0 12.1 0.0 Inocarpus fagifer 0.0 0.0 0.0 0.0 0.0 Ipomoea batatas 16.6 3.0 0.0 15.3 25.7 Musa sp.1 5.7 21.4 0.6 25.1 9.6 Musa sp.2 5.7 21.4 0.6 25.1 9.6 Piper methysticum 0.0 0.0 0.0 0.0 0.0 Solanum tuberosum 0.0 0.0 0.0 0.0 0.0 Spondias dulcis 0.0 83.3 0.0 0.0 0.0 Zingiber zerumbet 0.0 0.0 0.0 0.0 0.0 Average 5.1 13.5 0.2 6.8 2.5

Table 8.38 Yield ratios for archaeological species in all modern production systems (kg/time unit of labour)

Gadio Enga, New Guinea In the Gadio Enga production system, the species providing the highest output to input return ratio in terms of calories was Colocasia esculenta or the common taro (see Table 8.39). This cultigen had a ratio of 87,609.2kcal/instance of activity (ioa), while the next highest ratio was 196 that from Ipomoea batatas with 15,746.2kcal/ioa. Musa spp. had the highest ratios for arboreal species within the system, with 6507.7kcal/ioa. Another arboreal tree crop, Breadfruit or Artocarpus altilis had a ratio of 826.5kcal/ioa. Most of the Dioscorea yams ranked relatively low in comparison to these other cultigens, with ratios of 433.7kcal/ioa. The lowest overall calorific gain compared to labour investment was Zingiber zerumbet, which had a ratio of only 12.9kcal/ioa. These ratios also closely match the highest yield ratios for the Gadio Enga system.

The ratios for proteins, fats and carbohydrates follow much the same patterning and distribution as those for calories and overall yield. Clearly, the varying nutritional values for any of these species were insufficient to change the high-low distribution of ratios within the system. The distributions of values were largely dictated by the large differences between the yield and yield ratios for each species. Low ratios indicate that the production of species such as Dioscorea yams and Z. zerumbet is relatively inefficient, with low returns for time invested. These are gathered species and most likely require time for searching and harvesting that does not provide efficient returns compared to that produced from horticulture or even ‘silviculture’. In contrast the production techniques for species such as Colocasia esculenta and Ipomoea batatas within the Gadio Enga system are highly efficient, as significantly greater gains are made in terms of yield and associated nutritional values, for less time invested.

It is interesting to note that when these figures are compared with some other Highland PNG ethnographic datasets, the efficiency of some crop production is almost reversed. For example, within the Modopa (Waddell 1972) sweet potato (Ipomoea batatas) production has an output to input ratio of 8,075kcal/hr, while Dioscorea alata yams have a ratio of 10,258kcal/hr. These figures are lower and higher (respectively) than those calculated for the Gadio Enga, and reflect a primary focus on sweet potato production using large plano-convex mounding within open fields that are very rarely fallowed. Less time is invested in yam production within similar mounds in mixed gardens, but the calculated yield per hour is higher than that for sweet potatoes. In a Lowland PNG example, the Oriomo (Ohtsuka 1983) focus on sago production (2,000kcal/hr), with the horticultural production of other crops (980kcal/hr) primarily maintained to provide stability to the seasonal food supply. Chance shortages of sago was compensated by other staples from the gardens, both enriching diet and stabilising supply (Ohtsuka 1983:121). Wild plants only contributed seasonally to diet and in small amounts. Meat from hunted animals contributed around 67.1% of protein and 4.3% of energy, with an average of 25 minutes per day per capita dedicated to hunting compared to 118.8 mins for sago and 92.1 mins for horticulture (Ohtsuka 1983:119-18).

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Species Kcal/ioa Protein/ioa (g) Fat/ioa (g) Carbohydates/ioa (g) Alocasia macrorrhiza 0.0 0.0 0.0 0.0 Amorphophallus paeoniifolius 0.0 0.0 0.0 0.0 Artocarpus altilis 826.5 25.5 9.0 190.3 Colocasia esculenta 87609.2 1124.0 231.4 20894.0 Curcuma longa 0.0 0.0 0.0 0.0 Cyrtosperma merkusii 0.0 0.0 0.0 0.0 Dioscorea alata 433.7 5.9 0.4 91.0 Dioscorea bulbifera 433.7 5.9 0.4 91.0 Dioscorea esculenta 433.7 5.9 0.4 91.0 Dioscorea nummularia 433.7 5.9 0.4 91.0 Inocarpus fagifer 0.0 0.0 0.0 0.0 Ipomoea batatas 15746.2 265.2 16.6 3331.6 Musa sp.1 6507.7 65.9 17.2 1565.3 Musa sp.2 6507.7 65.9 17.2 1565.3 Piper methysticum 0.0 0.0 0.0 0.0 Solanum tuberosum 0.0 0.0 0.0 0.0 Spondias dulcis 0.0 0.0 0.0 0.0 Zingiber zerumbet 12.9 0.4 0.2 2.6

Table 8.39 Output to input ratios for archaeological species using Gadio Enga data

Bellona, Solomon Islands The highest yielding species for time invested in production in the Bellona system was Spondias dulcis (see Table 8.40). However, the nutritional output ratios suggest that this species would not necessarily have been preferentially cultivated or harvested over others within the system. This yield ratio is most likely the result of seasonal harvesting of this species from areas close to the village, so only a small amount of time is invested in searching, but high yields are gained for the time that is invested (Christensen 1975). While still high compared to many other species in the Bellona system, S. dulcis did not have the highest ratios for calories, proteins, fats or carbohydrates. The species with the highest ratio for calorific gain over time invested was in fact Cocos nucifera, with a ratio of 127,651.3kcal/hr, followed by S. dulcis with a ratio of 36,416.7kcal/hr. Artocarpus altilis also ranked relatively high with a ratio of 34,375kcal/hr. The lowest ranked species was Ipomoea batatas, which is interesting as this is a horticulturally produced cultigen. This species had a ratio of only 2,821.2kcal/hr— lower than the lowest ranked semi-cultivated species, Dioscorea bulbifera, which had a ratio of 3,102.7kcal/hr.

The ratios for protein follow much the same pattern as that observed within the distribution of calorific gain with the Bellona system. When the ratios according to fats are assessed, there are some slight differences. Cocos nucifera remains the highest yielding species with a ratio of 12960g/hr, followed by Spondias dulcis and Artocarpus altilis. However, within the lower ratios there are some changes. The two species with the lowest fat ratios are Amorphophallus paeoniifolius and D. bulbifera, two semi-cultivated species with ratios of only

198

1.8g/hr. Ipomoea batatas has a ratio of 3g/hr, which is just marginally greater than these two species.

Ratios according to carbohydrates produce further changes in the distribution of low-to- high ratios in the Bellona system. The highest gain in terms of carbohydrates is provided by A. altilis, with a ratio of 7916g/hr. Despite having an average carbohydrate content of 5-10/100g in comparison to other species within the system, C. nucifera still has the second highest ratio. This indicates that the techniques used for the subsistence production as well as export of this species were highly efficient, along with environmental suitability on the atoll (Christensen 1975). This species is followed by a number of horticultural root and tree crops that have high carbohydrate content, such as most Dioscorea yams, Musa spp., Colocasia esculenta and Alocasia macrorrrhiza. Ipomoea batatas had the lowest ratio of 596g/hr. When this figure is compared to the high carbohydrate content of this species (20.1/100g), it could be argued that the production of I. batatas within the Bellona system is not efficient in comparison with other horticultural crops. In other words, to gain calories, fats, protein and carbohydrates to contribute to daily diet with least time investment, I. batatas would not be the best selection in the recorded Bellona system. A more obvious choice would be either C. nucifera or A. altilis.

Species Kcal/hr Protein/hr (g) Fat/hr (g) Kg carbohydates/hr (g) Alocasia macrorrhiza 5803.6 106.8 8.2 1204.5 Amorphophallus paeoniifolius 3951.5 65.6 1.8 532.4 Artocarpus altilis 34375.0 1062.5 375.0 7916.7 Cocos nucifera 127651.3 1305.0 12960.1 3150.0 Colocasia esculenta 7254.5 93.1 19.2 1730.1 Curcuma longa 0.0 0.0 0.0 0.0 Cyrtosperma merkusii 0.0 0.0 0.0 0.0 Dioscorea alata 15189.4 261.4 10.2 2488.4 Dioscorea bulbifera 3102.7 56.8 1.8 739.4 Dioscorea esculenta 15265.5 224.6 22.8 2544.3 Dioscorea nummularia 13539.8 209.4 6.3 1690.2 Inocarpus fagifer 0.0 0.0 0.0 0.0 Ipomoea batatas 2821.2 47.5 3.0 596.9 Musa sp.1 24325.7 246.5 64.3 5851.0 Musa sp.2 24325.7 246.5 64.3 5851.0 Piper methysticum 0.0 0.0 0.0 0.0 Solanum tuberosum 0.0 0.0 0.0 0.0 Spondias dulcis 36416.7 666.7 250.0 3041.7 Zingiber zerumbet 0.0 0.0 0.0 0.0

Table 8.40 Output to input ratios for archaeological species using Bellona data

Anuta, Solomon Islands The recorded inputs and outputs for species in the Anutan system did not include many of the archaeologically identified species from Tongatapu. Therefore, there are fewer species to compare in terms of output to input ratios, with data available for only five crops (see Table 199

8.41). Of these, C. esculenta produced the highest ratios for most nutritional outputs, as well as yield. This indicates that of these species, C. esculenta. was produced with the highest efficiency in terms of returns for labour investment. This species had ratios of 1992.2kcal/hr, 8.5g/hr for protein, 5.3g/hr for fats, and 475.1g/hr in terms of carbohydrates. Likewise, Cyrtosperma merkusii, another aroid, had relatively high ratios of outputs to inputs with 1365 kcal/hr and 222.9g/hr of carbohydrates.

On Anuta, Yen (1973b) notes that the trees and perennial species within the agricultural system required the least attention during the cropping cycle, and in particular that bananas needed only limited weeding, thinning and the creation of windbreaks. It is then interesting to note that Musa spp. and Artocarpus altilis have lower yield and overall nutritional values than other staples.

There are some subtle differences in the efficiency of labour investment between two of these core staple aroids, Colocasia esculenta and Cyrtosperma merkusii. Cyrtosperma merkusii had relatively high ratios in terms of nutritional value apart from fats and protein, while C. esculenta had the lowest ratios for all nutritional values. This may be because Cyrtosperma is cultivated as a perennial species on Anuta, while a strict cycle of crop rotation is maintained for common taro and manioc (Manihot esculenta) which also involves harvesting for storage. The production of common taro is therefore more labour intensive, but apparently more efficient in terms of yield and nutritional returns.

Species Kcal/hr Protein/hr (g) Fat/hr (g) Carbohydates/hr (g) Alocasia macrorrhiza 0.0 0.0 0.0 0.0 Amorphophallus paeoniifolius 0.0 0.0 0.0 0.0 Artocarpus altilis 435.7 13.5 4.8 100.4 Cocos nucifera 0.0 0.0 0.0 0.0 Colocasia esculenta 1992.2 8.5 5.3 475.1 Curcuma longa 0.0 0.0 0.0 0.0 Cyrtosperma merkusii 1365.8 5.7 1.8 222.9 Dioscorea alata 0.0 0.0 0.0 0.0 Dioscorea bulbifera 0.0 0.0 0.0 0.0 Dioscorea esculenta 0.0 0.0 0.0 0.0 Dioscorea nummularia 0.0 0.0 0.0 0.0 Inocarpus fagifer 0.0 0.0 0.0 0.0 Ipomoea batatas 0.0 0.0 0.0 0.0 Musa sp.1 720.2 7.3 1.9 173.2 Musa sp.2 720.2 7.3 1.9 173.2 Piper methysticum 0.0 0.0 0.0 0.0 Solanum tuberosum 0.0 0.0 0.0 0.0 Spondias dulcis 0.0 0.0 0.0 0.0 Zingiber zerumbet 0.0 0.0 0.0 0.0

Table 8.41 Output to input ratios for archaeological species using Anutan data

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Tongatapu, Tongan Archipelago The most efficient horticultural species within the Tongan system in terms of yield and nutritional returns were Musa spp. (see Table 8.42). The production of these crops produced a yield to time investment ratio of 25.11kg/hr, and the difference in yield ratios compared to other species along with generally high nutritional values ensured that these crops also had high nutritional ratios. The calorific gain for labour investment for both Musa spp. was 32,751.1kcal/hr, while protein was 331.8g/hr, fats were 86.5g/hr and carbohydrates were 7,877g/hr. There is a significant difference between these species and those with the next highest ratios. Below Musa spp., the next highest ratios for calories, and carbohydrates were those from all yam species. In terms of fats Colocasia esculenta and Ipomoea batatas had higher ratios, while I. batatas also had a greater ratio for protein.

The least efficient species was consistently Alocasia macrorrhiza, which is not cultivated as commonly as other aroids such as C. esculenta or Xanthosoma sagittifolium on Tongatapu, according to the Agricultural Census (2001). Despite the size of the corms of this taro species, the flavour is considered less palatable, processing time is longer, and this species cannot be harvested as often as other varieties. It may also be the case that this aroid cannot be grown as efficiently as other aroids using dryland techniques. Depending upon the nutritional contribution needed by those cultivating and consuming these species, these differences in efficiency could impact the decision to cultivate more or less of a particular crop, or changes in the nature of labour investment through alterations to production techniques.

Species Kcal/hr Protein/hr (g) Fat/hr (g) Carbohydates/hr (g) Alocasia macrorrhiza 6674.4 122.8 9.4 1385.2 Amorphophallus paeoniifolius 0.0 0.0 0.0 0.0 Artocarpus altilis 0.0 0.0 0.0 0.0 Cocos nucifera 0.0 0.0 0.0 0.0 Colocasia esculenta 11857.6 152.1 31.3 2827.9 Curcuma longa 0.0 0.0 0.0 0.0 Cyrtosperma merkusii 0.0 0.0 0.0 0.0 Dioscorea alata 15545.3 212.5 15.2 3260.9 Dioscorea bulbifera 15545.3 212.5 15.2 3260.9 Dioscorea esculenta 15545.3 212.5 15.2 3260.9 Dioscorea nummularia 15545.3 212.5 15.2 3260.9 Inocarpus fagifer 0.0 0.0 0.0 0.0 Ipomoea batatas 14573.4 245.4 15.3 3083.4 Musa sp.1 32751.1 331.8 86.6 7877.6 Musa sp.2 32751.1 331.8 86.6 7877.6 Piper methysticum 0.0 0.0 0.0 0.0 Solanum tuberosum 0.0 0.0 0.0 0.0 Spondias dulcis 0.0 0.0 0.0 0.0 Zingiber zerumbet 0.0 0.0 0.0 0.0

Table 8.42 Output to input ratios for archaeological species using Tongan 2001 data 201

Ontong Java, Solomon Islands Similar to the Anutan system, Ontong Java had very little data for most of the species identified archaeologically on Tongatapu and therefore for this comparative exercise (see Table 8.43). Despite this, some assessment can be made from the existing data upon the efficiency of these crops in the Ontong Java production system. The species with the highest ratio for yield in terms of time investment was I. batatas, and this species also had the highest ratios for calories, protein, and carbohydrates. In terms of fats, the highest ratio was that from Cocos nucifera (78.5g/hr), which was significantly higher than the next highest ratio, derived from both Musa spp. (28.8g/hr). Both Colocasia esculenta and Cyrtosperma merkusii had roughly similar ratios when compared to other species. This is because these two crops are often grown together in freshwater swamps using similar techniques (Bayliss-Smith 1977:336). Although more land is dedicated to the production of Cyrtosperma, the yield ratios are similar and Colocasia has higher nutritional content in all values.

Species Kcal/hr Protein/hr (g) Fat/hr (g) Carbohydates/hr (g) Alocasia macrorrhiza 0.0 0.0 0.0 0.0 Amorphophallus paeoniifolius 0.0 0.0 0.0 0.0 Artocarpus altilis 0.0 0.0 0.0 0.0 Cocos nucifera 773.6 7.9 78.5 19.1 Colocasia esculenta 1214.7 15.6 3.2 289.7 Curcuma longa 0.0 0.0 0.0 0.0 Cyrtosperma merkusii 1927.0 8.1 2.5 314.5 Dioscorea alata 0.0 0.0 0.0 0.0 Dioscorea bulbifera 0.0 0.0 0.0 0.0 Dioscorea esculenta 0.0 0.0 0.0 0.0 Dioscorea nummularia 0.0 0.0 0.0 0.0 Inocarpus fagifer 0.0 0.0 0.0 0.0 Ipomoea batatas 24398.9 410.9 25.7 5162.3 Musa sp.1 10915.8 110.6 28.9 2625.6 Musa sp.2 10915.8 110.6 28.9 2625.6 Piper methysticum 0.0 0.0 0.0 0.0 Solanum tuberosum 0.0 0.0 0.0 0.0 Spondias dulcis 0.0 0.0 0.0 0.0 Zingiber zerumbet 0.0 0.0 0.0 0.0

Table 8.43 Output to input ratios for archaeological species using Ontong Javan data

System efficiency comparison and system classification When each of the archaeologically identified species are compared according to the ratio of outputs to inputs in terms of calories, proteins, fats and carbohydrates across each of the modern systems, it becomes clear that there are often significant differences (see Figures 8.22-8.25). As highlighted earlier in the analysis of these efficiency ratios within these systems individually, often the same patterning can be seen within all of the nutritional values. This patterning indicates that there are no significant differences in the ranking of various nutritional values for

202 species within these systems, and where there are changes in the ranking, the variances in values are not significant enough to overcome the difference in overall yield ratios between species. The consistency in patterning within output to input ratios can be seen within (Figures 8.22- 8.25).

Cocos nucifera or coconut provides an extreme example of these statistical differences in efficiency of production at species level. Within the Bellona system C. nucifera has a ratio of 12.7kcal per hour, while within the Ontong Javan production system it has 0.7kcal per hour. These differences are also reflected within other nutritional efficiency ratios for this species within these systems. Clearly there is difference in the production techniques used by these two systems. Ontong Java and Bellona both invested extra time into C. nucifera as part of a cash- cropping initiative to supplement subsistence (Bayliss-Smith 1977; Christensen 1975), but this labour input produced a much lower annual yield and associated calorific gain in the Ontong Java system.

In another example, Artocarpus altilis or breadfruit produced much higher yield and nutritional gain to lower labour inputs within the Bellona system (34,375kcal/hr, 1,062.5g/hr protein, 375g/hr fat, 7,916g/hr carbohydrates) than either the Gadio Enga (826.5kcal/instance of activity, 25.5g/ioa protein, 9g/ioa fat, 190.3g/ioa carbohydrates) or Anutan systems (435.7kcal/hr, 13.5g/hr protein, 4.8g/hr fat, 100.4g/hr carbohydrates). These three systems are based within very different environmental contexts. Bellona mostly consists of raised limestone, while Anuta is a small volcanic island and the Gadio live in the New Guinea highlands. These differing settings might partially explain the subsistence variation in nutritional efficiency ratios.

Similarly, Colocasia esculenta or the common taro produced a significantly higher yield within the Gadio Enga system (87,609.2kcal/instance of activity) using dryland techniques than any of the other systems, although this high ratio may be a result of the time unit used. Interestingly, the dryland production of taro on Tongatapu and Bellona also resulted in the generally higher output to input ratios of 11.827.6kcal/hr and 7,254.5kcal/hr, while wet techniques on Ontong Java and Anuta resulted in much lower ratios of 1214.7kcal/hr and 1992.2kcal/hr. In support of this trend, while only two dryland systems had recorded outputs for Alocasia macrorrhiza or the giant taro, the ratios from these were very similar. Within the Bellona system the ratio for this species was 5,803.6kcal/hr (106.7g/hr protein, 8.2g/hr fats, 1,204.5g/hr carbohydrates), while that in the Tongan system was 6,674.4kcal/hr (122.7g/hr protein, 9.4g/hr fats, 1,385.2g/hr carbohydrates) which is only marginally higher. Bayliss-Smith (1977) argues that the Ontong Java subsistence prior to the cash-cropping of copra was based on the production of taro, but the income generated by copra enabled the incorporation of new imported goods into production and consumption patterns. This change may have heavily impacted the efficiency of taro production on Ontong Java at the time this system was recorded.

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Again, there are also differences in the environmental contexts within which these systems are based, such as rainfall and geology, that will impact the efficiency of taro crop production.

The output to input ratios of these species were generally more similar across the different systems. This was often the case for many of the aroids other than C. esculenta, Dioscorea yams, Ipomoea batatas or sweet potato, and Musa spp. bananas or plantains. For example, I. batatas had output to input ratios of 15,746.2kcal/instance of activity (265.2g/ioa protein, 16.6g/ioa fat, 3331g/ioa carbohydrates) within the Gadio Enga system, 2,821.2kcal/hr (47g/hr protein, 2.96g/hr fats, 596.9g/hr carbohydrates) within Bellona, 14,573.4kcal/hr (245.5g/hr protein, 15.3g/hr fats, 3,063.4g/hr carbohydrates) within the Tongan system, and 24,398.9kcal/hr (410.9g/hr protein, 25.7g/hr fats, 5,162.3g/hr carbohydrates) within the Ontong Java system. The highest of these ratios were recorded within the Ontong Java system, and in this case I. batatas was grown within the village environs on a small scale along with several tree crops. The overall similarities in ratios, compared to others within these systems, could indicate that production techniques for sweet potato such as mounding, planting, harvesting and storage are relatively equal in terms of efficiency across different settings and scales. In another example, only two systems had recorded outputs for A. macrorrrhiza or the giant taro, but the ratios from these were very similar. Within the Bellona system the ratio for this species was 5,803.6kcal/hr while that in the Tongan system was 6,674.4kcal/hr— only marginally higher.

Overall, the greatest similarity among the ratios was between the Bellona and Tongan systems. These systems had ratios that varied only by 1000-3000kcal/hr for A. macrorrhiza, C. esculenta, Dioscorea alata, Dioscorea esculenta, and Dioscorea nummularia, and varied by 8000kcal for both Musa spp. These systems share a number of environmental similarities, such as a geological setting on a limestone raised island, and relatively consistent rainfall throughout the year, allowing the utilisation of dry production of taro. Both cultures also attach high social importance to the production of yams, and use similar propagation, planting and plant maintenance techniques such as trellising (Christensen 1975; Ministry of Agriculture and Forestry 2001).

The greatest difference in ratios was that between the Bellona and Anutan systems (see Figure 8.26). While the species within the Bellona system had an average ratio of 24,155.6kcal/hr (353.2g/hr protein, 1,060.5g/hr fats, 2872.1g/hr carbohydrates), the Anutan average was only 1,046.8kcal/hr (8.46g/hr protein, 3.14g/hr fats, 229g/hr carbohydrates). The Anutan system consistently had the lowest ratios for each species within this system, indicating that production techniques used within the mountain setting were generally inefficient in comparison with those used within other systems. The highest ratios of outputs to inputs within this system were those from Colocasia esculenta (taro) of 1992.2kcal/hr (8.5g/hr protein, 5.3g/hr fats, 475.1g/hr carbohydrates), suggesting that this was the crop that provided the

204 highest rate of return for lowest labour inputs. It is interesting that Yen (1973b: 139) asserts that the Anutan system was in his subjective opinion, “…one of the most intensive extant in the Pacific, despite the shortage of land and water, which could have conferred the potentiality for the well-known form of intensive agricultural production, irrigation farming of taro.” Yen (1973b) argued that the intensity of labour dedicated to agricultural production per hectare of land on Anuta in comparison to the swidden horticulture of the Hanunoo of Mindoro in the Philippines (3000/hectare versus 7000/hectare) supported this claim.

Based on the recorders’ own descriptions, these systems were classified as ranging from mixed to shifting or intensive agriculture. However, as argued by Dornestreich (1977:247-8), these terms are superficial and do not do justice to the full range of people’s actual subsistence behaviour. Instead, a subsistence typology should include information upon a) the environmental factors which substantially affect food getting, b) all the food-getting activities which compose the people’s subsistence system, and c) all the foods obtained by these activities in the form of a quantified account of food returns throughout an entire subsistence cycle (197:248). This comparative study has attempted to assess these Western Pacific example systems using these criteria, in order to provide a range of production systems within which the potential systems archaeobotanically identified on Tongatapu can be considered.

The intensity of labour investment in particular crops, using various production techniques that vary from cultivation through semi-cultivation to pure gathering of wild resources, has been compared to the overall yield and nutritional returns. There are of course limits to these data because of differences in what was deemed valuable by the recorders. Also, comparison of the intensity of labour between systems is problematic as each example varies in both scale and recorded time periods. Despite this, some descriptions can be made upon the efficiency of these systems in relation to the diversity of species exploited. Overall system efficiency was measured by dividing the total system efficiency by the number of species utilised within each system to create an average ratio of outputs to inputs. Patterning according to almost all nutritional values was the same, aside from carbohydrates (see Figure 8.26). Diversity of species is categorised as low (0-14 species), moderate (15-19 species), and high (20+ species). Yield ratio diversity is described as the difference between the average and highest value for all species, and categorised as low (0-10), moderate (16-30), high (31+). These systems can then be ranked in order of overall system efficiency and described as follows:

1. Bellona— high species diversity, insignificant nutritional diversity between groupings, significant labour diversity between groupings, high yield ratio diversity. 2. Tongatapu— moderate species diversity, insignificant nutritional diversity between groupings, insignificant labour diversity between groupings, moderate yield ratio diversity.

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3. Gadio Enga— high species diversity, insignificant nutritional diversity between groupings, significant labour diversity between groupings, high yield ratio diversity. 4. Ontong Java— low species diversity, insignificant nutritional diversity between groupings, insignificant labour diversity between groupings, moderate yield ratio diversity. 5. Anuta— moderate species diversity, insignificant nutritional diversity between groupings, significant labour diversity between groupings, low yield ratio diversity.

Figure 8.22 Output to input ratio comparison for archaeological species within each system in terms of calories

206

Figure 8.23 Output to input ratio comparison for archaeological species within each system in terms of protein

Figure 8.24 Output to input ratio comparison for archaeological species within each system in terms of fats (note vertical scale is logarithmic)

207

Figure 8.25 Output to input ratio comparison for archaeological species within each system in terms of carbohydrates

Figure 8.26 Comparison of average nutritional efficiency ratios for all systems (note vertical scale is logarithmic)

208

Comparison of prehistoric production systems The analysis of ethnographic examples from the Western Pacific has enabled the efficiency of crop production in different environmental settings to be assessed and discussed. These systems were ranked and described in terms of species and diversity of production technique (horticultural, semi-cultivated, gathered), as well as the efficiency of labour investment in terms of yield and nutritional returns. In this section the species identified within each of the archaeological sites of Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2) will be firstly compared to those species that have been ethnographically recorded within accounts of Tongan agriculture at the point of European contact, and are expected to be able to be identified archaeobotanically, in terms of overall nutritional value, labour and productivity. It was expected that these comparisons could facilitate discussion of the completeness of these systems, as well as system characterisation, when these are later compared to the Western Pacific example production systems during the second component of analysis.

Nutritional comparison of archaeological species Talasiu (TO-Mu-2) All of the species identified in the microbotanical and macrobotanical remains from Talasiu were compared and then ranked according to the same nutritional values discussed within the modern production systems (see Figure 8.27). The calories, protein, fats and carbohydrates within 100g of edible plant material from each archaeologically identified species were compared, and an overall rank given out of 14, where a low number equalled a high nutritional value. These species were then grouped into primary and supplementary species based on ethnographic data and the most common categorisation of these species within the example production systems.

Overall, Cocos nucifera and Inocarpus fagifer were ranked the highest and Piper methysticum was consistently ranked the lowest. Aroids such as Colocasia esculenta tended to be ranked high for overall nutrition figures (3rd), but varied from low to moderate ranking in terms of individual nutritional figures. Artocarpus altilis had very similar rankings to C. esculenta. The two remaining aroids, Cyrtosperma merkusii and Amorphophallus paeoniifolius, both ranked relatively high according to calories (5th and 4rd), and A. paeoniifolius also ranked high for protein (4th). With regard to fats, carbohydrates and overall nutritional figures these two species tended to rank low or moderately. Of the two yam species, Dioscorea alata generally ranks higher in all values than Dioscorea nummularia, which ranks very low (12th) according to overall nutrition. Both Musa spp. were ranked high for overall nutrition (5th) and carbohydrates (3rd), but ranked moderately according to calories, protein and fat. The two remaining supplementary species, Spondias dulcis and Zingiber sp., both ranked low for overall nutrition (13th and 12th) and most other values. The two exceptions to this pattern were that Zingiber sp. ranked high (4th) and S. dulcis ranked moderately (7th) for fats.

209

When these rankings are compared according to the groupings of primary and supplementary crops, some basic patterning can be determined. According to calories, the primary crops rank slightly higher, with an average ranking of 5.7 over the supplementary average of 8.3. The average rankings according to overall nutrition were almost exactly the same, with 5.9 for primary crops and 8.6 for supplementary. Comparison of protein rankings revealed that these two groups had almost equal averages of 7.4 within primary crops and 7.3 in the supplementary. In terms of rankings for fat content, the supplementary species actually tend to rank higher. These had an average of 6.3, while primary crops had an average of 7.6. Finally, according to carbohydrates the primary crops again rank higher with an average of 6, compared to 9.1 within the supplementary species. Overall these figures suggest that the primary crops tend to rank higher than supplementary, which confirms the fact that these species would have been given preference for cultivation.

Talasiu Nutritional Rankings 18 Primary crops Supplementary 16

10 Rank kcalories

Ranking 8 Rank protein

6 Rank Fat Rank carbohydrates 4 Rank nutrition

2 Primary crops Supplementary 0

Expon. (Rank nutrition)

Musa sp.2 Musa Musa sp.1 Musa

Cocos nuciferaCocos

Spondias dulcis

Dioscoreaalata

Artocarpusaltilis

Inocarpusfagifer

Pipermethysticum

Zingiberaceaetype

Dioscoreabulbifera

Colocasia esculenta

Cyrtosperma merkusii

cf. Dioscoreacf. nummularia Amorphophallus paeoniifolius Species

Figure 8.27 Nutritional comparison of species identified at Talasiu (TO-Mu-2)

To gauge whether Talasiu could represent a complete system, a list of species was developed from the ethnographic record of Tongan agriculture. This list was narrowed to include only those species that contained starch, endocarp, fruit or vegetative storage parenchyma that was likely to be processed and therefore enter the archaeological record at Talasiu (TO-Mu-2). The species that were identified in the archaeobotanical record at Talasiu were then compared nutritionally to these ethnographic species. When these two groups are graphed, the exponential trend lines for both archaeological and ethnographic species follow exactly the same curve (see Figure 8.28). When these two groups are compared statistically 210 using Student’s t-test, they cannot be differentiated, although the difference in averages is 43.06 (see Table 8.44). As to whether the species identified at Talasiu could represent a complete agricultural system, the question needs to be posed: do these archaeological species represent the best choices in terms of overall nutrition, and are they therefore more likely to be included in the diet at Talasiu? This comparison would suggest that they are generally equal to the ethnographic species in nutrition and so the results are inconclusive.

Talasiu and Expected Ethnographic Economic Species Nutritional Comparison 900 800 700 600 500 400 300 200 Archaeological

Total nutritionTotal figures (/100g) 100 Ethnographic 0 Expon. (Archaeological)

Expon. (Ethnographic)

Musa sp.2 Musa Musa sp.1 Musa

Cucumis melo

Cocos nuciferaCocos

Curcuma longaCurcuma

Puerarialobata

Dioscoreaalata

Spondias dulcis

Artocarpusaltilis

Ipomoea batatas Ipomoea

Inocarpus fagiferInocarpus

Morinda citrifolia

Benincasa hispida

Zingiberzerumbet

Pipermethysticum

Pandanus tectorius

Dioscoreabulbifera

Colocasia esculenta

Dioscoreaesculenta

Alocasia macrorrhiza

Syzygium malaccenseSyzygium

Cyrtosperma merkusii

Dioscoreapentaphylla

Dioscoreanummularia

Tacca leontopetaloides

Xanthosoma Xanthosoma sagittifolium Amorphophallus paeoniifolius Species

Figure 8.28 Nutritional comparison of species identified at Talasiu with expected ethnographic species.

Leka (J17) The ranking of species identified at Leka within both microbotanical and macrobotanical remains is compared in order to make suggestions about possible decision-making in terms of preferential cultivation and consumption. Leka has some additional species added to the provisions list compared to that of Talasiu, altering the rankings (see Figure 8.29). The highest ranked species in most values was Cocos nucifera, followed by Curcuma longa or turmeric, which is grouped here as a supplementary species according to ethnographic accounts of the use of this crop. Following these was Inocarpus fagifer, another supplementary crop, which ranked 3rd in all categories apart from protein and carbohydrates, for which this species ranked 2nd. Piper methysticum again ranked the lowest, followed closely by Spondias dulcis and Dioscorea nummularia. In most cases the rankings closely resemble those seen in the archaeological species identified at Talasiu, but would have lowered rank due to the high rank of C. longa. Of the yams, Dioscorea alata no longer ranks the highest due to the inclusion of D. esculenta which has higher nutritional rankings in all values apart from protein. A clear source of contamination was Solanum tuberosum, which was identified in the microbotanical remains. This species also ranks very low, aside from relatively high rank according to protein content (7th). 211

When these nutritional rankings are compared for the groupings of primary, supplementary and contamination species for Leka, some patterning can be discerned. In terms of calorific ranking, the primary crops rank slightly higher with an average of 7.8, while supplementary crops have an average of 8.3. Primary crops also rank slightly higher than supplementary species according to carbohydrates and overall nutrition, with average rankings of 7.7 and 7.8 over 8.6 and 8.5, accordingly. Interestingly, the supplementary crops rank higher than primary crops when the rankings for protein and fats are compared. In terms of protein, supplementary crops have an average ranking of 7.3, while primary crops have an average of 9.4. Similarly, the supplementary crops have an average fat ranking of 7, while primary crops have an average of 8.7. The contamination group ranks low compared to both groups in all categories apart from protein, for which this group has a rank of 6th.

Leka Nutritional Rankings 20 Primary crops Supplementary Contamination 18

10 Rank kcalories

Ranking Rank protein 8 Rank fat 6 Rank carbohydrates 4 Rank nutrition Primary crops 2 Supplementary 0 Contamination

Expon. (Rank nutrition)

Musa sp.1 Musa sp.2 Musa

Cucuma longa

Cocos nuciferaCocos

Spondias dulcis

Dioscoreaalata

Artocarpusaltilis

Inocarpusfagifer

Pipermethysticum

Dioscoreabulbifera

Colocasia esculenta

Dioscoreaesculenta

Solanum tuberosumSolanum

Cyrtosperma merkusii

Dioscoreanummularia Amorphophallus paeoniifolius Species

Figure 8.29 Nutritional comparison of species identified at Leka (J17)

As was the case with data from Talasiu, these species identified archaeologically at Leka were then compared nutritionally to those species that would be expected to be identified within the archaeobotanical record based on ethnographic accounts of Tongan agriculture (see Figure 8.30). These two groups were compared using the overall nutritional figures from 100g of edible plant material and graphed accordingly. The exponential trend lines for each were very similar, although there was some divergence, suggesting that archaeological species may have slightly lower overall nutritional value. However, when these two groups were compared statistically using Student’s t-test distribution, they could not be differentiated with any confidence (<50%). In fact, a comparison of the averages of these two groups provided a difference in mean of only 5.47. This comparison would suggest that the archaeological species

212 are almost exactly equal to the ethnographic species in nutrition and so the results are again inconclusive.

Leka and Expected Ethnographic Economic Species Nutritional Comparison 900 800 700 600 500 400 300

200 Archaeological Total nutritionTotal figures (/100g) 100 Ethnographic Expon. (Archaeological) 0

Expon. (Ethnographic)

Musa sp.1 Musa sp.2 Musa

Cucumis melo

Cocos nuciferaCocos

Curcuma longa Curcuma

Spondias dulcis

Puerarialobata

Dioscoreaalata

Ipomoea batatas Ipomoea

Artocarpusaltilis

Inocarpus fagiferInocarpus

Morinda citrifolia

Benincasa hispida

Zingiberzerumbet

Pipermethysticum

Pandanus tectorius

Dioscoreabulbifera

Colocasia esculenta

Dioscoreaesculenta

Solanum tuberosumSolanum

Alocasia macrorrhiza

Syzygium malaccenseSyzygium

Cyrtosperma merkusii

Dioscoreapentaphylla

Dioscoreanummularia

Tacca leontopetaloides

Xanthosoma Xanthosoma sagittifolium Amorphophallus paeoniifolius Species

Figure 8.30 Nutritional comparison of species identified at Leka with expected ethnographic species

Heketa (TO-Nt-2) The species identified using archaeobotanical techniques at Heketa were compared using the same nutritional values as those explored amongst species within the example production systems (see Figure 8.31). This list of species was markedly smaller than those from Talasiu or Leka, and so the rankings of species found at this site is different. As at Talasiu, the highest ranking species was Inocarpus fagifer, which ranked first according to all nutritional values, including overall nutritional figures. Piper methysticum also again ranked the lowest in all values apart from fat, which is ranked 3rd. Similarly, Spondias dulcis ranked very low in all values apart from fat, ranking 5th in this category. Of the aroids, Colocasia esculenta generally ranked the highest, with high to moderate rankings according to all values. The remaining aroids ranged from moderate to low ranking in all values, with a few exceptions. Alocasia macrorrhiza ranked high according to protein value (5th), while Cyrtosperma merkusii ranked high in calories (5th), and A. paeoniifolius ranked high in both calories and protein (4th). Ipomoea batatas is a possible modern or historic contaminant, but was included within the primary species for comparison, and ranks moderately according to protein (7th) and low in all other values. Only one Musa sp. was identified at the Heketa, and this ranked low according to carbohydrates (3rd), but moderately for calories, protein, fats and overall nutrition.

When these rankings are considered according to the grouping of species into primary, supplementary or contamination species, the nutritional value of core cultivated crops in 213 comparison with those that were most likely used on a more arbitrary basis can be assessed. In most cases, the difference in average rankings between these two groups is minimal. Primary crops were marginally higher ranked according to calories, with an average of 5.3 compared to the supplementary average of 5.75. The average rankings for protein were exactly the same for each group. Supplementary species ranked higher on average according to fat, with an average of 4.7 over an average of 5.6 in the primary crops. This was the only category in which supplementary species ranked higher than primary crops. The greatest difference between these two groups was observed within the rankings of carbohydrate content. Primary crops had an average ranking of 4.5, compared to an average of 7 within the supplementary species. Contamination species were not included in this comparison as it was deemed unnecessary.

Heketa Nutritional Ranking 14

12 Primary crops Supplementary

8 Rank kcalories

Ranking 6 Rank protein Rank Fat 4 Rank carbohydrates 2 Rank nutrition Primary crops 0 Supplementary

Musa sp.2 Musa Expon. (Rank nutrition)

Cocos nuciferaCocos

Spondias dulcis

Ipomoea batatas

Artocarpusaltilis

Inocarpusfagifer

Pipermethysticum

Colocasia esculenta

Alocasia macrorrhiza

Cyrtosperma merkusii Amorphophallus paeoniifolius Species

Figure 8.31 Nutritional comparison of species identified at Heketa (TO-Nt-2)

As a follow up to this nutritional analysis of archaeological species, these were then compared to species that could possibly also have contributed to Tongan diet and subsistence when Heketa was occupied (see Figure 8.32). The two groups again appeared to differ slightly according to overall nutritional figures for 100g of edible plant material, when the exponential trends are plotted on a bar graph distributing these species from highest value to lowest. However, a statistical comparison of these groups using Student’s t-test revealed that these groups cannot be differentiated according to nutrition (see Table 8.44, Table 8.45), despite an overall difference in values of 0.82 pooled standard deviations and a difference of 55.21 according to mean. These results have once again proven inconclusive in discerning whether the archaeological species could represent a complete system of nutritionally preferred species. 214

Heketa and Expected Ethnographic Economic Species Nutritional Comparison 900

800

700

600

500

400

300

200 Archaeological Total nutritionTotal figures (/100g) Ethnographic 100 Expon. (Archaeological) 0

Expon. (Ethnographic)

Musa sp.1 Musa sp.2 Musa

Cucumis melo

Cocos nuciferaCocos

Curcuma longa Curcuma

Puerarialobata

Dioscoreaalata

Spondias dulcis

Artocarpusaltilis

Ipomoea batatas Ipomoea

Inocarpusfagifer

Morinda citrifolia

Benincasa hispida

Zingiberzerumbet

Pipermethysticum

Pandanus tectorius

Dioscoreabulbifera

Colocasia esculenta

Dioscoreaesculenta

Alocasia macrorrhiza

Syzygium malaccenseSyzygium

Cyrtosperma merkusii

Dioscoreapentaphylla

Dioscoreanummularia

Saccharum officinarum

Tacca leontopetaloides

Xanthosoma Xanthosoma sagittifolium Amorphophallus paeoniifolius Species

Figure 8.32 Nutritional comparison of species identified at Heketa with expected ethnographic species

Nutrition (total figures/100g) Archaeological site Pooled standard deviation Pooled standard error Difference Confidence Mean difference Talasiu (archaeological vs ethnographic) 175.0 67.8 0.6 <50% 43.1 Leka (archaeological vs ethnographic) 173.7 67.3 0.1 <50% 5.5 Heketa (archaeological vs ethnographic) 180.1 67.4 0.8 50% 55.2

Table 8.44 Statistical comparison of nutritional value of archaeological and expected ethnographic species

Total nutritional figures (per 100g) Archaeological system Pooled standard deviation Pooled standard error Difference Confidence Mean difference Talasiu (primary crops vs supplementary species) 65.9 35.6 1.9 90% 68.9 Leka (primary crops vs supplementary species) 61.4 32.3 0.1 <50% 3.9 Heketa (primary crops vs supplementary species) 77.2 46.7 1.6 80% 73.2 All sites (primary crops vs supplementary species) 41.1 19.9 0.3 <50% 5.9

Table 8.45 Statistical comparison of nutritional value of species groups within archaeological systems at Talasiu, Leka and Heketa

Efficiency comparison of archaeological species and production systems The individual archaeological species from each of the sites of Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2) were combined into a list for each site, and then compared to the efficiency ratios provided by each of the modern Western Pacific systems. The comparisons enabled consideration of the efficiency (nutritional value of yield per time unit of labour

215 invested) of archaeological species in each site, and then an overall comparison of efficiency that may reflect the level of labour intensity within each time period represented by these sites. The species list from each site is here treated as a complete system, but as earlier data suggests, this is unlikely to be the case. Nevertheless, this analysis will speculate about the categorisation of these archaeological systems according to the spectrum provided within the modern production systems. These past systems will be placed in the context of current hypotheses upon the development of agriculture within Tongan prehistory in Chapter 9.

Inter-site comparison of species Each species in each of the Tongatapu sites has already been assessed according to output to input ratios that describe the efficiency of production techniques according to calories, protein, fats and carbohydrates. In this section, the combined species list from each site will be broken into the three groupings of primary crops, supplementary and contamination species and an efficiency comparison conducted according to calorific gain for time invested. The intention here is to model how these production systems may have functioned in the past, facilitating later discussion of preferences for cultivation. Clearly all of these species were utilised in some form, but which would have been staples? And how did these species enable the intensification of production to create surplus for the development of social hierarchy? To answer these questions, it is important to consider the ecology of agricultural systems in terms of context, inputs and feedbacks, and how these systems evolved over time.

When the species identified archaeobotanically in test units from Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2) are compared in terms of calorific efficiency, some suggestions can be made about crop preference (see Figures 8.33-8.35). Calorific efficiency ratios were selected as a focus; as previous assessments have suggested that most other nutritional value output to input ratios follow similar patterning. Of the 8 primary species and 6 supplementary species at Talasiu, Cocos nucifera has the highest average calorific efficiency ratio (64,210kcal/time unit) from all of the systems. This was followed by Spondias dulcis, primarily due to a lack of data from systems other than Bellona. Colocasia esculenta followed these species, with an average ratio of 21,500kcal/time unit and had the highest ratios of the aroids. Both Musa spp. had average ratios of 15,500kcal/time unit, just above Artocarpus altilis with an average of 11,700kcal/time. Of the yams, Dioscorea alata has the highest ratio average of 10,400kcal/time unit, marginally higher than Dioscorea nummularia which had an average of 9,800kcal/time unit. The only supplementary yam species, Dioscorea bulbifera, had a lower average of 6300kcal/time unit. The two remaining aroids, Amorphophallus paeoniifolius (3,900kcal/time unit) and Cyrtosperma merkusii (1.2kcal/time unit) both have very low averages compared to C. esculenta. Zingiber zerumbet had the lowest average compared to all other species, based on a single ratio of 1.3kcal/time unit from the Gadio Enga system. Two species, I. fagifer and P. methysticum had no data available and so were not included.

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Comparison of species at Leka revealed mostly similar distributions of average calorific efficiency ratios. These species included ten primary crops, five supplementary species, and one contaminant. Due to the inclusion of a few new species, some small ranking changes were observed. For example, D. alata does not have the highest average ratio amongst the Dioscorea yam species identified at Leka. Instead, D. esculenta, with a ratio of 10,400kcal/time unit has the highest average. This mirrors the nutritional figures discussed earlier in this section. Several species have no data from any of the systems, or are not particularly relevant as contaminants, and therefore were not compared. These include Curcuma longa, Inocarpus fagifer, Piper methysticum and Solanum tuberosum.

The list of species identified at Heketa is smaller than those from Talasiu and Leka, with only seven primary crops, six supplementary and no contamination species. Of these the top seven average ratios belong to species categorised here as primary crops. As at Talasiu and Leka, Cocos nucifera has the highest average ratio at Heketa. The inclusion of another aroid within this comparison changes the distribution amongst these taro species. Alocasia macrorrhiza had a higher average than both A. paeoniifolius and C. merkusii, with a ratio of 6,200kcal/time unit. Ipomoea batatas is possibly contamination within this site, but as explained earlier has not been classified as such here. This species has an average ratio that ranks between Musa sp.2 and A. altilis. Zingiber zerumbet again has the lowest average ratio of all species identified at Heketa. Three species had no data from any of the example systems and so could not be included in this comparison, including Curcuma longa, Inocarpus fagifer and Piper methysticum.

Statistically, the groupings of primary, supplementary and contamination species at Talasiu, Leka and Heketa were not markedly different according to calorific efficiency ratings. There was a difference in average of 7,100kcal/time unit at Talasiu between primary and supplementary groups, 1,900kcal/time unit at Leka, and 5,800kcal/time unit at Heketa. However, these groups could not be differentiated using Student’s t-test with any confidence above 50% based on the average ratios from all of the example production systems, which is unexpected considering the generally high averages of primary crops. When these ratios are then averaged themselves, these higher figures tend to become smoothed out over the number of species included in the comparison.

In light of this, it was deemed worthwhile to simply consider the ratios of primary to supplementary crops within each of these archaeobotanical assemblages. Talasiu has a ratio of 1.3:1, while Leka has a ratio of 2:1 and Heketa is 1.4:1. These indicate exploitation of a closer to equal range of economic and supplementary plant taxa at Talasiu, increased specialisation in primary crops at Leka, and some diversification again at Heketa.

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Figure 8.33 Comparison of calorific efficiency of archaeological species from Talasiu

Figure 8.34 Comparison of calorific efficiency of archaeological species from Leka

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Figure 8.35 Comparison of calorific efficiency of archaeological species from Heketa

Efficiency comparison of past production systems The overall system efficiency (in terms of calories) of these archaeological production systems was calculated in the same manner as for the modern systems and compared. Species from each of the sites of Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2) were combined into individual site lists, and then averaged according to the calorific efficiency values of each of the example production systems from Gadio Enga, Bellona, Tongatapu, Ontong Java and Anuta. This comparison enabled some overall comparison of the efficiency of the archaeological production systems from available data, if these systems are considered complete and resembled modern systems (see Figure 8.36). Clearly, the ratios for species within the Bellona production system were the most efficient in terms of calorific gain for time investment, while those from Anuta were the lowest. Therefore, if any of the three archaeological systems represented by archaeobotanical data at Talasiu, Leka or Heketa resembled the Bellona system, these would be more efficient in terms of calorific gain than if they were similar to the Anutan system.

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Figure 8.36 Modelled archaeological systems according to calorific efficiency values from modern systems

In light of these comparisons, Chapter 9 will consider the descriptions of each of the modern production systems in terms of species, nutritional, labour and yield diversity, as well as the environmental context and production techniques utilised. These descriptions will be used to predict the most appropriate models for archaeological production systems within this modern range. These models will also be considered in the context of current hypotheses about the antiquity of crop introductions and the subsequent development of agriculture in Tonga. It will be argued that, due to the lack of data available upon variables such as acreage dedicated to particular crops, scale of production, true labour investment, and dietary requirements of individuals, modelling archaeological systems against a range of known examples using Evolutionary and Human Ecological techniques can at least create new hypotheses upon the nature of agricultural systems in the past. It is important to note that social production will not be excluded from these models, as although this form of production is considered inefficient in terms of basic nutritional returns, this is still visible within this analysis and can be used to explain differences in productive efficiency between some species.

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Chapter 9 Discussion

This thesis sought to investigate the role of crops within colonising subsistence, as well as the links between the production systems in which these crops are cultivated and the development of social complexity in Tongan prehistory. With these aims in mind, the results of archaeobotanical investigations at Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2) are discussed here, focussing on modelling archaeological production systems. An agroecological approach to the analysis of agricultural systems was made in Chapter 8, weighing the nutritional costs and benefits of utilising particular plants in different environmental contexts. A scale was created for species diversity, nutritional value, labour investment and intensity, and yield diversity. Through this scale, the efficiency of both individual species and the system as a whole could be assessed to provide known parameters to model archaeological production systems. Chapter 8 provided the data and summarised patterning needed to be able to consider the most appropriate descriptions of systems in the past, as well as highlighting limitations. These models are placed in this chapter in the context of current hypotheses for Tongan and Western Polynesian agricultural development. Further, the antiquity of crop use will be considered in terms of the movement of people throughout the Western Pacific and also how crop use is tied to the development of social complexity in Tonga, culminating in the emergence of the state- level Tu’i Tonga paramount chiefdom.

Timing and nature of plant introductions into Tonga The chronology of crop introductions into Tonga and the development of production systems have been inferred through data from palaeoenvironmental research, linguistics and ethnographic studies (Burley and Connaughton 2007; Fall 2005, 2010; Fall and Drezner 2011, 2013; Kirch 1997; Maude 1965; Thaman 1976). In terms of economic species, there has been some conflict over the nature and timing of introductions due to differences in the information derived from oral traditions and that from scientific botanical, linguistic, genetic and archaeological research. Previous data and debates within Tonga and also the wider Pacific region will be considered here to address the first of the research questions of this thesis regarding the role of crops in the colonisation of the Pacific, alongside new data from the current research at Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2) (see Table 9.1). These data will be used to construct a new chronology for the timing of crop introductions into Tonga.

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Family Species Anacardiaceae Spondias dulcis Alocasia macrorrhiza Amorphophallus paeoniifolius Araceae Colocasia esculenta Cyrtosperma merkusii Arecaceae Cocos nucifera Convolvulaceae Ipomoea batatas Dioscorea alata Dioscorea bulbifera Dioscoreaceae Dioscorea esculenta Dioscorea nummularia Fabaceae Inocarpus fagifer Lecythidaceae Barringtonia asiatica Moraceae Artocarpus altilis Musaceae Musa spp. Piperaceae Piper methysticum Solanaceae Solanum tuberosum Curcuma longa Zingiberaceae Unknown

Table 9.1 List of all species identified archaeobotanically within this study

Spondias dulcis— Anacardiaceae Botanical research has suggested that Spondias dulcis, known commonly as Otaheite apple or vī, is probably native to the Indo-Malayan region, but was an ancient introduction in the Pacific as far east as the Marquesas (Whistler 2009). This species has not been recorded in any previous palaeoenvironmental research conducted in the Tongan archipelago (Fall 2005, 2010; Fall and Drezner 2011, 2013), or archaeobotanical research elsewhere in the Pacific, and therefore the relative timing of introduction to Tonga was not known. The presence of microbotanical remains of the species in the form of unmodified starch residues within well-dated deposits at Talasiu suggest that Spondias was introduced prior to 2750-2650 cal BP during the Lapita era. Spondias is not completely naturalised within island landscapes, arguably due to the large fruit which have no natural dispersers (Whistler 2009), and so would likely have been introduced as a cultivated supplementary fruit which was a source of food and medicine. Its use was historically documented by Cook (1785), La Billiardere (1793) and Waldegrave (1833). This species was also present throughout at Leka and Heketa.

Alocasia macrorrhiza— Araceae The giant taro or kape, is presumed to have been a prehistoric introduction into Tonga. This species is probably native to tropical Asia or New Guinea based on current distributions of wild species and subsequent cultivars which spread as far north as Hawaii and east to the Marquesas (Matthews 2014; Purseglove 1972; Whistler 2009). There is currently no palaeobotanical evidence for the introduction of A. macrorrhiza to any of the islands of Tonga. The earliest evidence from other locations within the Pacific document this aroid in the Solomon Islands at

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Kilu Cave by 28,000 BP (Loy et al. 1992), and north at Kosrae in Micronesia by around 1900 BP (Athens et al. 1996). Further east on Futuna, Piazza and Frimigacci (1991) documented the presence of pollen deriving from A. macrorrhiza at around 600 BP. The earliest evidence for the use of this cultigen in Tonga is dated to 800-600 cal BP from deposits within TP4 at Heketa. Ethnographic descriptions of the cultivation of this species in plantations at the time of European contact describe dryland production alongside yams, bananas, sweet potato and coconut (Cook in Beaglehole 1969; Cook 1785; Mariner 1817). Around eight varieties are recorded within Tonga that can be differentiated based on appearance (Maude 1965). Consumption of the stem was preferential to the root and leaves, which were often used to propagate new plants (Maude 1965).

Amorphophallus paeoniifolius— Araceae Amorphophallus paeoniifolius is one of the least commonly cultivated aroids in the Pacific and is known as elephant foot yam or more affectionately as the giant stink lily. The Tongan name for this species is teve. Literature describes a larger reliance upon this species in the recent past, but it is now utilised primarily as a famine food and is largely semi-cultivated or naturalised in secondary forest and plantations (Purseglove 1972; Whistler 2009). Records of this species within Tongan history are limited to one possible mention by Cook (1785:332), where he describes “…a large root called ‘kape’, one not unlike our white potatoes called ‘mawhawha’; the taro, or coccos of other places, and another called ‘jeejee’”. This could refer to the teve. A later study noted the presence of Amorphophallus in bush allotments, but described these plants as uncultivated and, according to informants, eaten only in times of famine (Thaman 1976). Archaeobotanical investigations in this study have identified the presence of A. paeoniifolius at all three sites, indicating that this species was introduced early by 2750-2650 cal BP during the later stages of Lapita settlement on Tongatapu.

Colocasia esculenta— Araceae The common taro is arguably one of the most versatile cultigens incorporated into the production systems of the Pacific region. It can be grown in a variety of different climatic and geological contexts, using wet irrigation or dryland shifting cultivation techniques (Kirch 1994; Leach 1999; Spriggs 1996, 2002; Yen 1973a). It was thought native to Southeast Asia (Whistler 2009), but has been found within archaeological deposits in the Solomon Islands dated to 28,000 cal BP (Loy et al. 1992), at Niah Cave in Malaysia by 23,850-23,020 cal BP (Paz and Barton 2007) and in Papua New Guinea by 10,220-6440 cal BP at Kuk Swamp (Denham et al. 2003), indicating that the wild progenitor of this species may have had a wider distribution than was thought (Matthews 2014). The taxonomy and domestication of this species is also complicated by the distribution of Colocasia formosana in Southeast Asia, a wild naturally dispersed taro that is phenotypically close to C. esculenta (Matthews et al. 2015). Prehistoric movement of C. esculenta into Remote Oceania is attributed to the Lapita Cultural Complex as

223 far eastwards as Western Polynesia and Polynesians are most likely responsible for initial migration and subsequent adaptation of this crop to the diverse island landscapes of Central and Eastern Polynesia.

The earliest evidence of Colocasia near Western Polynesia is at Bourewa in Fiji at 3050-2500 BP through starch grain, raphide and xylem tissue identification (Horrocks and Nunn 2007). Starch residues on pottery from Upolu, Samoa have been confirmed as common taro within deposits dated to the Late Lapita occupation of the region by 2750 BP (Crowther 2009). Starch, pollen and parenchyma of taro has been identified in many post-Lapita or late Holocene deposits in Rapanui (Cummings 1998; Horrocks and Wozniak 2008; Horrocks et al. 2012 ), New Caledonia (Horrocks, Grant-Mackie and Matissoo-Smith 2008), Kosrae (Athens et al. 1996), Mangaia (Kirch et al. 1995), Hawaii (Allen 1981, 1984), the Marquesas (Allen and Ussher 2013), Pitcairn (Horrocks and Weisler 2006) and New Zealand (Horrocks and Barber 2005; Horrocks and Lawlor 2006; Horrocks, Smith, Nichol, Shane and Jackman 2008; Horrocks, Smith, Nichol and Wallace 2008; Horrocks et al. 2004, 2007). Previous palaeoenvironmental research in Tonga has identified pollen from Colocasia within swamp cores from Vava’u and Eua dated to around 2600 BP (Fall 2005, 2010).In the current research parenchyma and starch grains of Colocasia esculenta were identified in deposits from Talasiu, Leka and Heketa. Results indicate that this crop was at least introduced into the Tongan archipelago by 2750-2650 cal BP through Late Lapita migration and was utilised throughout Tongan prehistory.

Cyrtosperma merkusii— Araceae Native to Melanesia, it has been argued that Cyrtosperma merkusii or the giant swamp taro may have been one of the first species cultivated in Remote Oceania by the Lapita settlers of the region. The brackish conditions in beach back-swamps near many early Lapita settlements would have enabled the initial cultivation of saline-resistant crops such as Cyrtosperma prior to the establishment of more labour-intensive irrigation required for other aroids such as Colocasia or Alocasia (Kirch and Lepofsky 1993; Yen 1973a, 1982, 1993). On atolls Cyrtosperma is grown within various sized pits dug to access the freshwater aquifer (Weisler 1999), and so this is another possible cultivation technique that would have enabled early settlers to survive on raised limestone islands such as Tongatapu where traditional irrigation of common taro (Colocasia) was impossible. The earliest evidence of Cyrtosperma within the Pacific is from pollen dated to around 4500 BP in mangrove cores taken from Ngerchau region of Palau (Athens and Ward 2001), although this may be due to natural distribution (Athens and Stevenson 2012). Within Remote Oceania, starch, raphides and xylem tissues of giant swamp taro was identified within Late Lapita-associated deposits from Urupiv in Vanuatu, dated to around 2700 BP (Horrocks and Bedford 2004; Horrocks et al. 2014). Other evidence of cultivation has been found in more recent deposits on Kosrae dated to 1997-1350 BP (Athens et 224 al. 1996), the Pitcairn Group after 950 BP (Hather and Weisler 2000), and parenchyma on Mangaia from 788-331 BP (Kirch et al. 1995). Microbotanical evidence for Cyrtosperma at Talasiu, Leka and Heketa, represents the first recorded identifications of this species in the Tonga. The presence of this taxa within deposits at Talasiu, a late Lapita-associated site, indicate that this cultigen may indeed have been grown using the same pit agricultural techniques observed in Micronesia until more intensive dryland techniques for other aroids and yams were fully developed and this became a minor staple.

Cocos nucifera— Arecaceae Cocos nucifera, known commonly as coconut or niu in Tonga, was often present prior to human arrival due to dispersal of the large floating fruit on ocean currents, surviving up to 110 days before husks become waterlogged (Gunn et al. 2011;Whistler 2009). It is undoubtedly one of the most important cultivated species in the Pacific region, providing a source of water, food, fuel, and numerous household items such as food storage vessels and thatch (Gunn et al. 2011). The nutritional analysis of Cocos within this study points to the high calorific, protein, and fat content of the mature meat, especially compared to other common cultigens. The only core nutrient that this species lacks is carbohydrates, which is often provided by multi-cropping cultivation with starchy root, tuber and tree crops in plantations. There have been some attempts to associate the cultivation of coconut with Lapita settlement in Island Melanesia at Arawe (Matthews and Gosden 1997), which may have actually been the result of beach drift, and more confidently on Mussau from 3200-2000 BP (Kirch 1987, 1988, 1989). In Tonga, this species is documented within pollen records prior to Lapita arrival but an increase in pollen quantities around 2600 BP could indicate the cultivation of Cocos for food after this time (Fall 2005, 2010; Fall and Drezner 2011, 2013). The identification of charred endocarp deriving from Cocos nucifera at Talasiu confirms this association, and continued cultivation is proven by the presence of macrobotanical remains in deposits at Leka and Heketa, and also through many ethnographic records (Beaglehole and Beaglehole 1941; Cook 1785; Gifford 1929; Orlebar 1830; La Billardiere 1800; La Perouse 1799; Mariner in Martin 1991; Waldegrave 1833; Wilson 1797).

Ipomoea batatas— Convolvulaceae The cultivation of sweet potato (Ipomoea batatas), or kumala, is an important dryland crop in Tonga. Ipomoea was listed as a major crop by Maude (1965), and was recorded within several agricultural censuses (Ministry of Agriculture and Forestry 1985, 1994, 2001). The timing for the introduction of this South American cultivar into Tonga is disputed. Two possibilities are enabled within the ‘tripartite hypothesis’ proposed by Barrau (1957) and developed by Yen (1974), Green (2005) and Clarke (2009). The first is that this species was an early Polynesian introduction of plants within the ‘Kumara’ line from Central and Eastern Polynesia (Roullier et al. 2013). The second possibility is that Ipomoea was a later introduction during the 18th or 19th

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Centuries by either Polynesians or Europeans, supported by Kirch’s research on Niuatoputapu (1978, 1990). Genetic research does not provide much clarity in Western Polynesia (Denham 2013; Roullier et al. 2013). Early historic accounts clearly document the presence of this cultigen as early as the late 18th century (La Billardiere 1793) and into the 19th century onwards (Gifford 1929; Beaglehole and Beaglehole 1941; Waldegrave 1833). It is therefore most likely that it was in fact a Polynesian import, and is supported by archaeobotanical evidence from Eastern and Central Polynesia which documents the cultivation of Ipomoea on Mangaia from around 950 BP (Hather and Kirch 1991), in the Marquesas by 750-350 BP (Allen and Ussher 2013), Hawaii at 650-325 BP (Horrocks and Rechtman 2009), Rapanui by 650-150 BP (Cummings 1998; Horrocks and Wozniak 2008) and in New Zealand to the south by 704-550 cal BP (Horrocks et al. 2007). Considering these dates and the maritime connections of the Tongan state, it is not unlikely then that Ipomoea batatas was present within the archaeological system at Heketa from at least 600 cal BP, and is not a modern contaminant. Further, the archaeological starch was also compared to that of beach morning glory (Ipomoea pes-caprae) which grows near the sea, but did not match the morphology of that species.

Dioscorea spp.— Dioscoreaceae Yams have played a very important role in Tongan agricultural and cultural practices throughout prehistory. Three main named types are recognised ‘ufi tokamu’a (early yam), ‘ufi tokamu’i (late yam) and ‘ufi lei (sweet yam), but there are many variants within each type. The early yam and late yam have been botanically identified both as Dioscorea alata, while the sweet yam is Dioscorea esculenta. During his third voyage Cook described these two species as “The roots are yams of which are two sorts, one black, and so large, that it often weighs 20 to 30 pounds; the other white, and long, seldom weighing a pound” (1785:331-2). In 1643 Tasman traded European goods for provisions for the ship from Tongatapu that included yams (Tasman 1776). Other species that have been ethnographically recorded are Dioscorea bulbifera which is naturalised and the aerial bulbils are often used as a famine food (Waldegrave 1833; Whistler 2009), Dioscorea nummularia and Dioscorea pentaphylla, although these are less commonly cultivated. Early ethnographers and missionaries commented on the importance of yams within Tongan festivals such as the ‘inasi or first fruits festival, the tau tau ceremony, and pongipongi feasting (Cook in Beaglehole 1969; Cook 1785; Gifford 1929; Mariner in Martin 1991). Others noted the arrangement of yam crops within plantations, methods for cultivation, and annual crop cycling. Cook (1785) specifically commented on the distinct layout and functional divisions within individual plantations, as well as the differences between the produce reserved for the chiefly elite and the commoners.

None of these Dioscorea spp. have been identified within palaeoenvironmental or archaeological deposits within the Tongan archipelago prior to this study. The recovery of D. alata, D. nummularia and D. bulbifera at Talasiu therefore provides the earliest evidence for the 226 introduction of this genus into Tonga, and data from Leka indicates continued cultivation into the Formative Period. The presence of D. esculenta in deposits dated to 800-600 cal BP at Leka suggests that this may have been a later introduction, or is perhaps due to preservation bias. In any case, the arrival of yams can only be securely dated to after this time within the Formative Period. Elsewhere in the Pacific, the use of Dioscorea yams has been documented as early as around 40,000 BP at Niah Cave in Malaysia (Paz and Barton 2007), and 10,220-9100 BP at Kuk Swamp in PNG (Denham 2007). Lapita-associated use has been indicated by the recovery of microbotanical remains identified as D. esculenta from Fiji and Vanuatu (Horrocks and Nunn 2007; Horrocks et al. 2013), and D. nummularia and D. pentaphylla also from Vanuatu (Horrocks and Bedford 2010). Other evidence from Central and Eastern Polynesia support the early transportation of these crops further into Remote Oceania. In fact, yams were transported to each corner of the Polynesian Triangle as far as Hawaii in the north (Allen 1984), Rapanui to the east (Horrocks and Wozniak 2008; Horrocks et al. 2012), and New Zealand in the south (Horrocks and Barber 2005; Horrocks, Smith, Nichol, Shane and Jackman 2008; Horrocks, Smith, Nichol and Wallace 2008). These place the findings of this study within the broader Western Pacific context, and indicate that yams played an important role in Lapita and post- Lapita subsistence.

Inocarpus fagifer— Fabaceae The Tahitian chestnut or ifi, is cultivated as a supplementary species in plantations or found naturalised in secondary forest. It is believed to have originated in the Indo-Malayan region and has a current distribution as far east as the Marquesas (Walter and Sam 2002; Whistler 2009). Early explorers and ethnographers recorded the presence of ifi in Tongan subsistence. One of the earliest was La Billardiere who described the placement of and consumption of this species. He wrote: “The sugar canes we saw there, were planted at a pretty considerable distance from each other, under the shade of inocarpus edulis [polynesian chestnut], the fruit of which these people roast and eat, its flavour much resembling that of the chestnut.”(La Billardiere 1793:366). Two other early ethnographers who recorded the cultivation of this species were Gifford (1929) and Waldegrave (1833).

The earliest recorded evidence for the consumption of Inocarpus within the Pacific region is from the Lapita-associated site of Mussau, where pericarp dated to 3200-2000 BP was recovered (Kirch 1987, 1988, 1989). Very little further evidence for the prehistoric use of this species has been found in archaeobotanical studies. Huebert (2014) recovered small quantities of Inocarpus wood charcoal on Nuku Hiva in the Marquesas, associated with deposits dated after the 17th century. In Micronesia, Inocarpus was identified on Kosrae by Athens and others (1996) through parenchyma relatively dated to around 1900 BP. The identification of starch from Inocarpus fagifer within this study is the first microbotanical evidence for the cultivation of this food, and the first evidence for the introduction of this species into Tonga by 2750-2650 227 cal BP. This species was present throughout the strata at Talasiu, Leka and Heketa indicating that the nature of starch morphology and chemistry for this species facilitates high rates of preservation. It is also possible that this supplementary species was more heavily relied upon in the past.

Barringtonia asiatica— Lechythidaceae The fish-poison tree or futu is not an introduced species to Tonga, having a native distribution from Madagascar through to the Marquesas in the east in coastal stands of vegetation (Walter and Sam 2002; Whistler 2009). It is not consumed, but rather the seed within the endocarp is removed and grated to produce a pulp. The pulp is then mixed with sand and wrapped in leaves before being placed in shallow areas of lagoons where there is little water circulation and fish are forced to breach the surface to escape the effects of the poison. Fish are then caught easily in nets (Whistler 2009). The species has only been recorded rarely in archaeobotanical assemblages in the Pacific, and mostly within relatively recent deposits after 500 BP (Huebert 2014). Pollen records from Vava’u in Tonga have confirmed the presence of this species from around 2600 BP in sampled marine clays along with Rhizophora mangroves, Thespesia populnea, Guettarda, Cocos nucifera and Pandanus tectorius (Fall 2005). This palaeoenvironmental evidence supports the identification of Barringtonia asiatica within deposits at Talasiu and Leka from 2750-2650 cal BP. It is unlikely that starch grains of the seed of Barringtonia entered the archaeological record at Talasiu through natural aeolian or hydraulic processes, as the seed would need to somehow be exposed, dehydrate and then be abraded enough to release reserve starch grains that were blown or washed into deposits. This suggests that the species was being used for cultural and subsistence purposes (fishing) during late Lapita occupation at least through to the Formative Period.

Artocarpus altilis— Moraceae Artocarpus altilis is a domesticated variety of breadfruit or mei deriving from A. camansi (breadnut) which is both wild and cultivated in New Guinea (Jones et al. 2012; Kennedy and Clarke 2004). There is also hybridisation with Artocarpus mariennensis, endemic within Micronesia (Ragone and Raynor 2009), cultivars from which may have also been distributed across the Micronesian-Polynesian divide. Domestication processes, timing and spread of these varieties is complex, and not yet fully understood. In Remote Oceania, Artocarpus altilis and A. altilis X A. mariennensis hybrids are divided into two types, seeded and seedless cultivars. Seedless varieties are taxonomically identified as A. altilis, and tend to be cultivated for carbohydrates contained within the starchy fruit. Seeded varieties are either identified as A. altilis or hybrids of A. altilis and A. mariennensis, and are often cultivated for both the flesh and the seeds that are also a source of starch. Despite some varieties producing seeds, A. altilis is not propagated through seeds, but rather from root and shoot cuttings from which seedlings are produced (Ragone 2006). The geographic and chronological details of the development and

228 spread of breadfruit remain to be fully investigated. (Jones et al. 2012; Kennedy and Clarke 2002)

Although the species does not play a prominent role within subsistence today on Tongatapu, early ethnographic evidence suggests that this crop may have been much more popular within plantations in the past (Beaglehole and Beaglehole 1941; Cook 1785; Gifford 1929; Orlebar 1830; La Billardiere 1800; La Perouse 1799; Mariner in Martin 1991; Tasman 1776; Waldegrave 1833; Wilson 1797). Dutch explorers La Maire in 1616 (1967 reprint) and Tasman in 1643 (1776) observed that provisions they traded for included breadfruit, however it was not as commonly grown as in the Society Islands. Cook (1785:272) describes the breadfruit as the first food articles that his crew could purchase when they first arrived, and the trees themselves were interspersed with little order alongside coconut and close to villages. La Perouse in the 1780’s (1799:170) also described Tongatapu as having “…no trees; but such as like the cocoa, breadfruit, that affords them subsistence”. Archaeobotanical evidence for the cultivation of breadfruit has been found within a number of post-Lapita sites in Polynesia and Micronesia. The starch, pollen, wood or exocarp parenchyma of this crop has been identified in Kosrae (Athens et al. 1996), Society Islands (Kahn and Ragone 2013; Orliac 1997), Marquesas (Allen and Ussher 2013; Huebert 2014), and possibly on Futuna (Piazza and Frimigacci 1991). Prior to this study, there had been no palaeoenvironmental or archaeological evidence for the cultivation of breadfruit in Fiji-West Polynesian prehistory. The recovery of starch and possibly the storage parenchyma of the fruit of A. altilis at Talasiu indicate the species may have been one of the initial Lapita crop introductions to Tonga. Further, the species was likely cultivated throughout prehistory, as evidenced by small numbers of preserved starch grains at both Leka and Heketa.

Musa spp.— Musaceae Cultivated forms of the genus Musa are a complex, confused and confusing group, of great importance in the Pacific and elsewhere. At least three distinctive lineages of cultivated bananas have been described for Oceania, all of them with at least partial New Guinea ancestry. Although biomolecular evidence has shown the complexities of these lineages, they are yet to be distinguished in the archaeobotany of Remote Oceania (Kennedy 2008). They can be difficult to differentiate taxonomically, and have therefore been simply classified as Musa spp. within the reference collection and archaeobotanical identifications. The first domestication of the Musa genus probably occurred within the Pacific region. The earliest evidence for possible cultivation derives from Kuk Swamp in Papua New Guinea where phytoliths extracted from archaeological deposits date to 10,200-6400 cal BP (Denham et al. 2003, 2004; Fullagar et al. 2006; Wilson 1985). There is also convincing evidence that Musa spp. were transported into Remote Oceania by Lapita times. Starch and phytoliths from these crops have been identified within Lapita associated deposits in the Reef Islands of the Solomons (Crowther 2009), and on 229

Urupiv and Matilau in Vanuatu (Horrocks and Bedford 2010; Horrocks et al. 2009). Furthermore, Musa phytoliths, pollen, endocarp and vegetative storage parenchyma have been recovered in Eastern and Central Polynesia from a number of locations (Allen 1984; Horrocks and Rechtman 2009; Horrocks et al. 2012; Kirch et al. 1995; McAllister 1933; Piazza and Frimigacci 1991). The data from Talasiu, Leka and Heketa on Tongatapu fits well within this chronology and distribution. But which particular lineages within the Musa genus were cultivated by the Lapita settlers of Tongatapu and the Western Pacific, and transported throughout the rest of Remote Oceania, remain to be established. Given the great diversity of Oceanian cultivated bananas, it is quite likely that multiple movements , replacements and accretions have contributed to this set of vital arboreal crops that are low maintenance but also high yielding and are still cultivated today.

Piper methysticum— Piperaceae The kava root is used throughout Polynesia and Micronesia as a relaxant often during traditional festivals and rituals, and for medicinal purposes. It is thought to be native to Vanuatu where it was first utilised and domesticated as a male clone of the endemic species Piper wichmannii (Whistler 2009) before being transported east to West Polynesia and north to Micronesia. It is solely propagated by hand through root and stem cuttings as populations outside of Vanuatu are all male, which gives strength to the hypothesis that this species was anthropogenically- introduced to all other islands in the Pacific where it is currently grown (Balick and Lee 2009). Currently, there is no direct evidence for the timing of the domestication of kava within Vanuatu, and the movement east. However, a number of studies support the hypothesis that this may have occurred during Lapita settlement and interaction through analysis of ceramics that resemble kava bowls and cups (Green 1974; Palmer 1968), linguistics (Geraghty 1983), chemistry and biology of kava varieties (Lebot et al. 1997), and skeletal analysis of temporomandibular joint (TMJ) degeneration that can be tied to kava preparation (Visser 1994). Archaeobotanical evidence for kava cultivation and consumption has been recovered from late prehistoric deposits in the Marquesas from 550-350 BP (Allen and Ussher 2013), and Hawaii (Allen 1984) although a fossilised stem fragment belonging to a member of the Piperaceae family has been identified in deposits associated with Lapita artefacts and dates to 3600-2400 BP (Kirch 1989).

Evidence from early ethnographic accounts point to the prominent role that this sedative played in Tongan culture. Cook’s (1785) surgeon, Mr Anderson, commented that kava was commonly planted about their houses, and that the root was the only part dug up for ceremonies. Servants were observed breaking the roots into pieces, scraping the dirt off with a shell, and preparing the mix (1785:318). Cook (1785:282) himself noted that on Tongatapu there was “…hardly anywhere without the kava plant, from which they make their favourite liquor.” La Billardiere’s crew (1793:339) were treated to a dinner of yams roasted in the fire embers and 230 plantains, which La Billardiere assumed were to take the heat from the stomach produced by the ‘intoxicating liquor’ of kava. Furthermore, he describes “...its stalk, frequently bigger than the thumb, is tolerably straight, and requires no support. They cut off several pieces in the spaces between the knots, and they made us a present of them, informing us, that they set them in the ground, in order to propagate the plant.” During the 1920’s Gifford (1929) observed that most festivals and ceremonies involved the presentation of both food and kava to higher chiefs such as the Tu’i Tonga or Tu’i Kanokupolu. One of the most detailed accounts of kava consumption was published by Collocott (1927), where he describes the etiquette and placement of titled individuals during kava ceremonies within a ‘kava-ring’. The careful distribution of roots is also described in detail along with the methods for preparing the brew, with some comments on modern adaptations of traditional practices. At least seven different types of kava are described in terms of root size and how these are brought to these ceremonies.

Accounts and archaeological evidence from elsewhere in the Pacific are important for assessing the nature of kava consumption in the past, and the timing of its introduction and cultivation in Tonga. The microbotanical evidence from Talasiu (TO-Mu-2) suggests that this crop was probably imported to Tonga during initial colonisation, but possibly in the latter part of this period. This data ties in with current evidence suggesting late Lapita transportation of kava out of northern Vanuatu during a period of interaction that facilitated the development of the Proto-Polynesian language (Green 1966; Kirch 1989; Lebot et al. 1997; Visser 1994). The presence of starch deriving from Piper methysticum within well-dated deposits at Leka (J17) and Heketa (TO-NT-2), indicates the continued cultivation and incremental inclusion of this crop within Tongan social practices. It is likely that the use of kava became tied to the development of social hierarchy within Tongan prehistory, and was used to validate positions within this hierarchy, as observed by Collocott (1927) and Gifford (1929) even after European contact.

Curcuma longa and Zingiber spp.— Zingiberaceae Members of the ginger family have played important roles in Tonga and the wider Pacific as food items, but also for medicinal and domestic purposes. Most are thought to have been first cultivated in Southeast Asia, but are now widely distributed throughout Melanesia, Polynesia and Micronesia (Purseglove 1972; Whistler 2009). Curcuma longa or turmeric (ango) is cultivated primarily as a dye for colouring tapa cloth or for body decoration. The methods for processing the harvested roots varies throughout the Pacific, but generally involves some washing, grating, straining, cooking and drying (Whistler 2009). There are only a small number of ethnographic accounts that note the cultivation of turmeric in Tonga, including Cook (1785), and Gifford (1929). Recent studies of Tongan agricultural production have noted the rare presence of turmeric within allotments, often in a wild state, but protected due to the value of the rhizome for the preparation of dye (Thaman 1976). It may be that this crop is preferentially 231 cultivated closer to the village or within house gardens, rather than bush allotments. Other varieties of ginger are not mentioned within early ethnographic accounts pertinent to Tonga, but Zingiber zerumbet was recorded by Whistler (2009) during botanical survey. This particular species of Zingiber was used as a shampoo, to add scent to tapa, and the sap could also be drunk. There have been no recovered archaeobotanical remains of any plants belonging to Zingiberaceae in the Pacific prior to this study. Here the vegetative storage parenchyma of the rhizome of an unknown species within the Zingiberaceae family was identified in late Lapita- associated deposits at Talasiu, and the starch of Curcuma longa was extracted from sediments at Leka that can be dated to within the Formative Period of Tongan prehistory from 1300-1000 cal BP. This evidence points to the use of members of this family within Tongan subsistence from at least 2750-2650 cal BP, and continued archaeobotanical investigations on Tongatapu may further elucidate the chronology of use for gingers in Tonga.

Modelling archaeological production systems Each of these plant introductions in Tongan prehistory were either cultivated or utilised as supplementary species in tropical production systems. The nutritional values and efficiency (inputs vs. outputs) of each of these taxa extracted from the archaeological sites included in this research (Talasiu, Leka and Heketa) were compared in Chapter 8. Here, these were combined to characterise and discuss each of the archaeological production systems. These modelled systems were used to address each of the two key research questions regarding the role of plants in the subsistence of colonising populations in Tonga, and within the development of the social complexity seen through evolution of the maritime chiefdom or state. Critically, it would appear that modelled decreased system nutritional efficiency is linked to increased specialisation in primary crops, observed through the ratios of numbers of primary to supplementary crops within each of these archaeobotanical assemblages. Discussion will focus on how people may have been cultivating or exploiting plants found in the archaeobotanical record, based on the context of modern production systems, and also how changing system efficiency over time can be linked to species diversification or specialisation.

Feasibility of modelling Two main issues concern the feasibility of modelling archaeological systems, based on both the data available and the approach used in this thesis research. It is important to address whether the list of species identified in various micro- and macrobotanical remains at each site included in this study might represent a ‘complete’ production system. Due to limitations in the faunal data at all three sites, a full comparison with the protein dietary components was deemed inappropriate. Further, without knowing the specific dietary requirements of the populations occupying each of these sites in the past and the contribution of other food sources, it cannot be calculated whether plant or animal species could have provided sufficient nutritional and energetic benefits. These dietary requirements depend on the age, gender, and lifestyle of each 232 member of the population, and therefore cannot be gauged from the limited data available at Talasiu, Leka and Heketa. When these archaeological plant species are compared with other ethnographically recorded species in accounts of Tongan agriculture, there is no statistical difference in overall nutrition between the groups. This remains the case even when the list restricted to only those species that are likely to be processed during food preparation and then preserved in these archaeological deposits. Future analysis of modern soil samples from agricultural systems to gauge the presence and quantity of plant microfossil and macrobotanical material would enable greater accuracy in the analysis of plant inputs to outputs from archaeological samples.

For each of the archaeological sites, statistical comparisons were made using overall nutritional figures (combination of calories, fat, protein and carbohydrates within 100g of edible material), and compared using Student’s two-sided t test and distributions. None of these were statistically different with over 95% confidence and so the null hypothesis that these could have been from the same population was unable to be rejected. These results are interpreted here to suggest that there was no major nutritional advantage to be gained by choosing any of the ethnographic species in addition to those found within the archaeological assemblages. So it is plausible that these archaeological species could be viewed as composing ‘complete’ systems, based purely on overall nutrition. Comparison of yield ratios may reveal a different patterning, but this data is variable and is not available for all species. It would also be unlikely that all of the ethnographically recorded species were introduced during initial settlement and were therefore present throughout the Tongan sequence. The implication of this being that species were most likely introduced gradually over time, and could feasibly be represented by the ranges of species found archaeobotanically.

The second issue concerns the approach taken to modelling the archaeological systems. It is argued here that an agroecological approach allows alternative hypotheses to be explored through basic comparison of nutritional ranking, with overall system efficiency measured in terms of the nutritional or energetic value of estimated yield (outputs) per unit of time invested in labour (inputs). Using criteria that have been developed to describe a modern range of ethnographic production systems from the Western Pacific, these archaeological systems were characterised and modelled in terms of overall calorific efficiency (as assessment of individual nutritional values revealed that patterning of system efficiency was generally equal) according to predicted labour investment. This overall system nutritional efficiency was based on the labour and yield data from the modern system that most closely resembled each archaeological system, based on these characterisations.

The specific features of modern and archaeological systems explore a range of options for labour investment and associated yield and nutritional return ratios that attempt to avoid the

233 issues of geographic and temporal scale, breaking overall efficiency down into outputs per man- hour for ease of comparison. Of course, this analysis does not provide any measure of social or cultural benefits that require consideration of efficiency at the level of individuals. It is a purely nutritional and energetic approach to productivity that explores decision-making through the cultivation capacity of plants identified archaeologically based on a range of recorded and described Pacific production systems.

Expected modelling outcomes Based on current hypotheses for island colonisation and settlement, it was expected that this modelling would indicate a pattern of developing complexity and intensity within production systems over time from Talasiu (TO-Mu-2) (2750-2650 cal BP) through to Heketa (TO-Nt-2) (800-600 cal BP). After weighing the arguments for Lapita subsistence, it is clear that the ‘strandlooper’ or solely marine-based forager-type hypothesis (Best 1984; Groube 1971) is unlikely in light of current isotopic, faunal, and botanical evidence (Horrocks and Bedford 2004, 2010; Horrocks and Nunn 2007; Horrocks et al. 2009, 2014; Kinaston et al. 2013; Kinaston, Bedford, Richards, Hawkins, Gray, Jaouen, Valentin and Buckley 2014; Kinaston, Buckley, Valentin, Bedford, Spriggs, Hawkins and Herrscher 2014; Valentin et al. 2010). Initial dispersal of populations may not have involved the introduction of plants, but this quickly changed within one or two generations. If Lapita subsistence strategies did involve some element of terrestrial production, the question remains as to the degree of reliance on imported plants over any endemic or native species, and how these plants were cultivated. The argument over mental templates within colonising populations involving knowledge of wet taro production using pondfield construction and irrigation has been discussed partly in Chapter 2 (Bellwood 2005, 2011; Kirch and Lepofsky 1993; Spriggs 1990, 1996, 2003; Yen 1973a). These arguments for and against independent development of pondfield irrigation are mostly irrelevant when discussing horticultural systems on Tongatapu. This raised limestone island is not suitable for wet taro production and extensive dryland techniques were utilised instead. However, this argument is worth considering in relation to the types of crops that may have been brought to the archipelago and experimented with. These agricultural techniques may also have been brought to Tonga and Samoa before moving out into Central and Eastern Polynesia after a 1500-year pause that saw the transition to Ancestral Polynesian Society. It seems unlikely that independent innovation of pondfield irrigation occurred over and over throughout the Pacific, and yet current archaeobotanical and archaeological evidence does not suggest that colonising populations utilised these techniques until much later in historical sequences. Instead it has been argued that early Lapita migration into Western Polynesia was initially supported by ‘broad- spectrum’ subsistence strategies, as defined by Ingold (2000) and Kennedy and Clark (2004) utilising both cultivation systems and marine-based technology (Hunt 1980; Kirch 1978:12). With regard to Tonga specifically, Burley (1998; Burley and Dickinson 2001) agrees, promoting Kirch’s (1994) refined definition of a middle ground position, whereby agricultural activities 234 were initially of secondary importance and possibly limited to low energy swidden-type cultivation.

The arguments generally depend on the timing and speed of the migration from Near Oceania into Remote Oceania. New data from Bayesian analysis of radiocarbon dates from the Bismarcks, Fiji and Vanuatu suggests that there was only a pause of around 130-290 years from 3360 to 3240BP in this eastern end of Near Oceania before colonisation began in Island Melanesia and Western Polynesia by 3250-2900 BP (Denham et al 2012:44; Sheppard 2011). New data from Tonga indicates the archipelago was also first colonised relatively quickly around 2900-2800 (Burley et al. 2015). This relatively fast migration might suggest that agricultural techniques such as pondfield irrigation would not necessarily provide an advantage within subsistence practices at this early stage of colonisation. Causal factors such as population growth and environmental change are unlikely to have forced the adoption of labor-intensive techniques within such a short time frame.

One ecological approach attempted to operationalise migration episodes through the Ideal Free Distribution (IFD) concept. Kennett, Anderson and Winterhalder (2006) used this technique to account for modes of subsistence, new habitat suitability, and population density in predicting migratory behaviour in the colonisation of East Polynesia. The concept predicted a ‘leap-frogging’ type of behaviour, with periodic episodes of migration followed by population growth. Modes of subsistence are seen as the key variable influencing population growth and environmental or habitat variability. For example, low level and later intensive food production contributed to faster decreases in habitat suitability through environmental degradation, but concurrently increased the overall carrying capacity of many remote island habitats (Kennett et al. 2006:268). Overall the archaeological data from Oceania is argued to be consistent with the predictions of the IFD model, with the long interval between the initial settlement of Near Oceania at around 35,000 BP and movement in to Remote Oceania at around 3300 BP matching the curve for foragers living on large islands (Kennett et al. 2006:285). Further, the rapid migration into Remote Oceania after the colonisation of Western Polynesia is consistent with the predicted curves for food production on small islands, while the intensification of these systems can be viewed as an implication of the 1500-year pause here before later migration into East Polynesia.

Considering these arguments, it was hypothesised that the Late Lapita production system at Talasiu, several hundred years after initial settlement, would most likely have resembled ‘broad-spectrum’ subsistence. This system would be characterised by high species diversity including a number of both supplementary native and introduced species as well as known Lapita cultivated cultigens such as a range of aroids, yams and tree crops. Production techniques would focus predominantly on multi-cropping annual and perennial species that

235 would provide short-term returns for labour investment. Arboriculture would probably still play a small role in this system and would have low efficiency in terms of yield and nutritional benefits from labour investment. The comparative ethnographic system that best matches this description would be the Gadio Enga production system, which utilises plants within a number of agro-ecosystems that range from fully cultivated or managed to just simply gathered native or naturalised species.

Looking beyond Late Lapita subsistence at Talasiu, the timing of occupation at Leka (J17) represents settlement of the area during the Formative Period from around 1300-1000 cal BP. Due to a lack of archaeological or archaeobotanical data, nothing is known about the role of plants within subsistence strategies at this time. There is no evidence that the integrated maritime Tu’i Tonga chiefdom had yet emerged, and the monumental architecture associated with this chiefdom had not yet been constructed. The use of the phrase ‘the Dark Age’ for this period by Janet Davidson (1979:94-5) is still relatively accurate, as the disappearance of pottery after 1500 BP has meant that little is known about the nature of local cultural developments until around 1000 BP (Burley 1998). Sites become both difficult to find and define. Limited material remains from excavated sites suggest relative continuity between the Plainware and Formative Periods; however, faunal data from Niuatoputapu indicate less reliance on marine resources and increased production of pig (Kirch 1988). Data that has been correlated to the end of the Formative Period point to an expansion of earthen burial mound construction, probably associated with population growth (Burley 1998; Spenneman 1989).

Archaeobotanical remains from occupation at Leka reflect subsistence technology influenced by concepts brought with the Lapita colonisers or through later migration and interaction, and also by local development. It is clear that the fragile ecological composition of islands, in conjunction with their insularity, makes them environmentally dynamic and susceptible to any change. In an ongoing dialectic between environmental-human-climatic factors, modes of subsistence are both created and restricted by environmental and social factors within this dynamic ecological setting. Leppard (2014:10) has argued, however, that environmental thresholds may not be the most valuable concept for visualising carrying capacity early on in island sequences, but rather culturally-created thresholds that may have been crossed. Pressure on food resources required innovative ways of alleviation, such as expansion to new niches, intensification of food production systems or new modes of distributing resources (Leppard 2014:10). Modes of distribution are argued by many Pacific researchers to also include exchange networks within and between island groups (Green 1987; Kirch 1987, 1997; Summerhayes 2000; Terrell 1989). Unfortunately limited archaeological evidence during this post-Lapita period in Tonga makes the association of subsistence practices with cultural development difficult. Interaction was frequent in early and late phases of Tongan prehistory, suggesting that this also the case in the ‘Dark Age’. It can only be assumed that any changes 236 that occurred paved the way for the subsequent development of the classic Tu’i Tonga chiefdom, and multi-tiered social hierarchy (Davidson 1979). All of these environmental and cultural factors, as well as those transported mental templates for production strategies already discussed, would have characterised the form and flexibility of production at Leka.

With these factors in mind, it is likely that subsistence during the post-Lapita Formative Period was characterised by continued experimentation with introduced cultigens, some intensification of production to cope with environmental, social or demographic factors, and further exploitation of natural resources. Studies of environmental change on Tongatapu have pointed to a gradual drawdown of sea level from the mid-Holocene hydro-isostatic highstand that would have drastically changed the shoreline of the Fanga’ Uta Lagoon (Dickinson 2001, 2007). The changing lagoon environment impacted the extraction of resources from this inshore ecosystem, and is argued to have resulted in a significant decrease in the size and variability of faunal marine species, particularly shellfish, through over-exploitation (Grono 2012). This change was particularly clear in cultural deposits from Leka TP4 (Grono 2012). Clearly, there was a change in the focus of subsistence from the marine to the terrestrial environment based on previous evidence from occupation at Leka and other sites dated to the Formative era in the Tongan archipelago.

The background to subsistence and cultural development at Leka would suggest that the production system represented by archaeobotanical data from this site would have high species diversity through continued periods of interaction and migration. This expansion of the economic plant repertoire on Tongatapu should also correlate with significant nutritional diversity between primary and supplementary crops. Continued experimentation with production techniques suitable for the Tongan environment involving both new and established cultigens would have resulted in increased overall efficiency in terms of energy and nutritional returns, but would also have increased labour diversity between staple and other crops. For these reasons, it was expected within this study that the archaeological production system evidenced by cultural deposits at Leka would most likely resemble the Bellona system. This system was characterised by shifting dryland cultivation with some intensive production for the cash- cropping of copra, supplemented by the exploitation of supplementary species.

The final phase of Tongan prehistory, represented by archaeobotanical data from this research, is the transition to a centralised maritime chiefdom which integrated the entirety of the Tongan archipelago by at least 500 BP. Two deposits from test units at Heketa dated to around 800-600 cal BP and 600 cal BP, and fall into a period characterised by the development of monumental architecture at Heketa as an early centre for chiefly political control (Clark and Reepmeyer 2014; Spenneman 2002). At this later end of the Tongan sequence, subsistence is argued elsewhere to have become more intensive or extensive to develop surplus to support the

237 increasing socio-political complexity of the Tongan chiefdom (Aswani and Graves 1998; Clark et al. 2008; Poulsen 1983, 1987; Spenneman 1989). Other proximal causes have also been emphasised, such as environmental and demographic factors and warfare, but primacy has been given to extensive and labour-intensive dryland food production strategies (Kirch 1984). Modelling of agricultural dryland production and increasing population by both Green (1973:69-73) and Kirch (1984:222) indicates that all of the arable land on Tongatapu could have come under production by AD 1000 or possibly earlier. The timing of this ceiling on the availability of land is argued to coincide with the amalgamation of local control into a central polity on Tongatapu, and the beginning of a period of maritime expansion that extended Tongan influence as far afield as Fiji, Samoa, Niue and Futuna (Aswani and Graves 1998:144). In his interpretive model, Kirch (1984) elaborated on Green’s (1973) initial hypothesis of land pressure triggering population control measures, arguing that population growth and competition for land resulted in a series of regional chiefdoms that constantly assimilated smaller groups to increase overall control and ended with the formation of a central polity on Tongatapu. This was based on Carneiro’s (1970) hypothesis for state formation.

Correlation between the development of monumental architecture on Tonga and emerging social complexity has also been drawn by a number of researchers. McKern (1929) was the first to study this evolutionary confluence, and argued that the organisation of manpower required to construct these stone platforms was the logical result of centralised political and social power held by the Tu’i Tonga. Others have since expanded this hypothesis. For example, Clark and others (2008) have argued that the change in form from non-sepulchral architecture to large burial platforms at around AD 1450 coincided with the political integration of the chiefdom under the 24th Tu’i Tonga, Kau’ulufonua 1, and reorganisation of titles into the sacred (Tu’i Tonga) and the secular (Tu’i Ha’atakalaua). The transition to sacred chief began with the move of power from Heketa to Lapaha by the 12th Tui Tonga, Talatama, but was intensified by this further stratification. Oral traditions also link the 24th Tu’i Tonga to an increase in territorial expansion to Uvea, Futuna and Samoa. Clark and others (2008) speculate that at this crucial stage in the Tongan chronology an emphasis on the semi-divine status of the chiefly lineages resulted in an increase in social complexity through further stratification of the political hierarchy, but also increased the risk of chiefs distancing themselves from practical government and control. It can be postulated that the segment of the population providing the manpower required for construction may have been unable to produce their own food, and so surplus would also have been needed to feed these workers. Redistribution of food for this purpose may have resulted in the emergence of the tribute system, although this is difficult to prove.

More recently, discussion has centred on the nature and scale of interaction within Western Polynesia and how this influenced developing social complexity. There is some 238 evidence for early interaction within this region during Lapita settlement and the development of Ancestral Polynesian Society or more classic Western Polynesian culture (Best 1984; Burley et al. 2011; Clark et al. 2014). Later in Tongan prehistory, further development of trade networks may have both extended and reinforced Tongan influence within Fiji and Samoa during the development of the maritime state (Clark et al. 2014). These long-distance political and economic exchanges within the second millennium AD have been traced through the sourcing of lithic material from state and pre-state contexts on Tongatapu, and post-ceramic contexts from Samoa. Results indicated a non-local long-distance source for 66% of analysed stone artefacts from Tongatapu, but predominantly local sources for the Samoan material. Interarchipelago interaction was clearly a feature of early occupation on Tongatapu that intensified with the development of stratified society. It was argued from this data that an important consequence of social complexity was the establishment of new types of specialised sites for the transmission of people, plants and materials (Clark et al. 2014).

The connection between expansion of horticultural production and social complexity is supported by early ethnographic accounts of intensive cultivation practices and tribute systems evident during the yearly pattern of festivals such as the ‘inasi or first fruits festival (Gifford 1929; Mariner in Martin 1991). Intensification of production arguably enabled the development of political economies within other islands and archipelagos, such as Hawaii (Earle 1980, 1991, 2012; Graves et al. 2011; Kirch et al. 2012; Kirch 1994; Ladefoged and Graves 2008, 2011; McCoy and Graves 2012), Futuna (Kirch 1976, 1984, 1994), and the Marquesas (Addison 2006; Allen and Addison 2002; Allen 2010; Earle 1993; Rolett 1998). The transition from broad- spectrum subsistence strategies to more labor-intensive dryland agriculture was not a simple process. It required decisions and political control of land to change cultivation and processing techniques which increase the availability of essentially lower-ranked and less efficient resources. This was likely to be a response to the decreased availability of higher-ranked resources in terms of nutrition and energy efficiency, sourced through hunting or gathering. However, the pathway to agricultural development through agroforestry, arboriculture, irrigated vegeculture, or rain-fed vegeculture depends on both environmental constraints and cultural factors. Understanding the mechanisms of change during this transition through the intentionality within varying levels of decision-making is critical, and is the means by which these can be linked to the current chronology of migration and socio-political development.

With these discussions in mind, it is expected that the archaeological production system at Heketa, if complete, would most closely resemble the analysed Anutan production system. It is likely that plant production at this point focussed primarily on dryland crops such as taro and yams that would produce the largest yields within short periods of time, but not necessarily the greatest nutritional returns. Overall system efficiency would be lower due to the decrease in species diversity, and the utilisation of labour-intensive methods for crop production through 239 shortening of fallow lengths and reduction of production areas, after an initial period of expansion. Labour diversity between groupings of plants (primary or supplementary) would therefore also have become significant.

In summary, the archaeological plant production systems represented by archaeobotanical remains from Talasiu (TO-Mu-2), Leka (J17), and the Heketa (TO-Nt-2) are expected to have been similar to those from Gadio Enga, Bellona and Anuta respectively. These expectations were based purely on weighing current arguments of the nature and role of subsistence within Late Lapita, Formative Period, and the monumental state. The expectations are compared with the outcomes of modelling archaeological systems using an agroecological approach to view these systems in terms of species, nutritional and possible yield diversity, as well as labour efficiency considering outputs for crops, groups (primary or supplementary) and the overall system.

Modelling Talasiu (TO-Mu-2) The dense and expansive midden at the site of Talasiu (TO-Mu-2) represents a well-dated snapshot of subsistence in the Late Lapita period through faunal, floral and artefactual material. AMS dates from test unit TP2 have pinned the occupation of Talasiu to around 2750-2650 cal BP based on short-lived charred endocarp extracted from throughout the unit. The narrow range of all dates from these deposits suggests that the midden at Talasiu represents settlement on the shoreline of the Fanga ‘Uta Lagoon for around 100 years or less. Talasiu is therefore not the earliest dated site on Tongatapu, as the site of Nukuleka closer to the entrance of the lagoon has been U/th dated to 2838±8 cal BP (Burley 2001a,b; Burley et al. 2015), but archaeobotanical data from Talasiu can provide information about the early post-settlement phase of Tongan prehistory. As the Lapita cultural complex evolved into Polynesian society, whose descendents would later colonise Central and Eastern Polynesia, production systems played a major role in facilitating this process. This vital transitional phase is reflected within the data from Talasiu, and can be modelled using an ecological approach to production systems that considers the range of utilised species, and the nutritional efficiency of these that would influence decision- making regarding the types of production techniques engaged in cultivation, management or opportunistic gathering practices.

The range of species identified within archaeobotanical remains from Talasiu TP2 represents a mix of arboreal species and root and tuber crops from a number of different families (see Table 9.2). Only one of these taxa is considered endemic or native to Tongatapu, and the remainder must therefore have been introduced by Lapita colonisers or their descendents. Cocos nucifera has been found within pre-human deposits in pollen cores from Avai’o’vuna Swamp on Vavau during a marine high-stand dated to 4500-2600BP (Fall 2010), and likely arrived through the flotation of fruit upon marine currents. All but two of the

240 identified taxa are generally accepted to have been early prehistoric introductions into Tonga. One exception is Ipomoea batatas, known more commonly as the sweet potato, which is a known later introduction from South America via Eastern Polynesia that is suspected by some researchers to even be a historic introduction (Kirch 1978, 1990), and was therefore classified as modern contamination within this assemblage. The other is Piper methysticum or kava, which some oral traditions in Tonga suggest was a late prehistoric export from Vanuatu, but has not so far been corroborated with linguistic or genetic data. Considering this it has been included within the list of species, primarily to keep an open mind about this crop in terms of the timing of its introduction and the role this species may have played within production systems in the past. There are also a number of species that have not previously been identified within Lapita- associated deposits in Melanesia or Western Polynesia. These include the aroid Amorphophallus paeoniifolius, the yam Dioscorea bulbifera, Inocarpus fagifer (Tahitian chestnut), Barringtonia asiatica (Fish-poison tree), Artocarpus altilis (breadfruit), and Piper methysticum or kava.

Family Species Anacardiaceae Spondias dulcis Amorphophallus paeoniifolius Araceae Colocasia esculenta Cyrtosperma merkusii Arecaceae Cocos nucifera Dioscorea alata Dioscoreaceae Dioscorea bulbifera Dioscorea nummularia Fabaceae Inocarpus fagifer Lecythidaceae Barringtonia asiatica Moraceae Artocarpus altilis Musaceae Musa spp. Piperaceae Piper methysticum Zingiberaceae Unknown

Table 9.2 Identified families and species within archaeobotanical remains from Talasiu (TO-Mu-2)

Using the criteria derived from the analysis of production systems from the Western Pacific, the archaeological production system at Talasiu (at least that based on those species identified archaeologically) can be described and compared to this range of ethnographic examples. Compared to this range of systems, Talasiu has moderate species diversity (15 species), and has insignificant nutritional diversity between groupings of ‘primary crops’ and ‘supplementary species’ (90% confidence). The labour and yield diversity of species within the Talasiu system cannot be known. However, when these species are compared using the figures from all of the example systems, a range of possibilities can be gauged. In the case of Talasiu, the average yield ratios (see Table 9.3) within the archaeological system can range from low (0- 10) to high (31+) diversity, depending on the figures used. Despite large differences between the average labour inputs for each grouping when these are modelled using the figures from the 241 example systems (see Table 9.4), these are not deemed to be statistically significant differences when calculated using Student’s t-test (95% confidence or over based on the number of compared samples). When modelled on the Bellona Island data, labour investment between the two groups could be differentiated by 2.04 standard errors, but based on the number of samples (degrees of freedom) the probability of samples or species being from the same group is 20%. This probability does not allow rejection of the null hypothesis that these two groups could belong to the same population based on mean difference (see Table 9.5). Under these conditions the Talasiu system could be characterised as:

 Talasiu— moderate species diversity, insignificant nutritional diversity between groupings, insignificant labour diversity according to groupings, low-to-high yield ratio diversity.

Based on this, the Talasiu system most likely resembled the production system recorded on Tongatapu in 2001 in the Agricultural Census of that year. If this is the case then the average calorific efficiency of this system is around 20666kcal/hr, with a total calorific efficiency of 123996kcal/hr (see Figure 8.36). When it is considered that the data from the 2001 Tongatapu system was based on an intensive cash-cropping economy, this characterisation is unexpected. It was expected that the Talasiu system would more closely resemble the Gadio Enga production system, which was described as ‘mixed’ or ‘diverse’ by Dornstreich (1977) involving similar labour investment into each species within a range of exploited agro-ecosystems, if late Lapita subsistence fell somewhere between a ‘strandlooper’ and ‘transported landscape’ economy. Using the comparative analysis carried out during this study, the Gadio Enga system was characterised as having high species diversity, insignificant nutritional diversity between groupings, significant labour diversity between groupings and moderate yield ratio diversity. Clearly these do not match the description of Talasiu outlined here, and this discrepancy will be discussed at the end of this section.

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Species Gadio Enga Bellona Is Anuta Tongatapu Ontong Java Amorphophallus paeoniifolius 0.0 2.9 0.0 0.0 0.0 Artocarpus altilis 1.0 41.7 0.1 0.0 0.0 Cocos nucifera 0.0 30.0 0.0 0.0 0.5 Colocasia esculenta 66.1 5.5 0.0 8.9 0.9 Cyrtosperma merkusii 0.0 0.0 0.5 0.0 1.6 Dioscorea alata 0.3 12.7 0.0 12.1 0.0 Dioscorea bulbifera 0.3 2.9 0.0 12.1 0.0 Dioscorea nummularia 0.3 12.7 0.0 12.1 0.0 Inocarpus fagifer 0.0 0.0 0.0 0.0 0.0 Musa sp.1 5.7 21.4 2.5 25.1 9.6 Musa sp.2 5.7 21.4 2.5 25.1 9.6 Piper methysticum 0.0 0.0 0.0 0.0 0.0 Spondias dulcis 0.0 83.3 0.0 0.0 0.0 Zingiber zerumbet 0.0 0.0 0.0 0.0 0.0 Average 5.7 16.8 0.4 6.8 1.6

Table 9.3 Yield ratios for species identified at Talasiu modelled using comparative systems

Gadio Enga Bellona Is Anuta Tongatapu Ontong Java Grouping Species (instances of activity) (hours/yr) (hrs/37 days) (hrs/yr) (hrs/yr) Primary crops Artocarpus altilis 10.3 60.0 62.6 115.2 183.0 Cocos nucifera nd 3333.3 30.3 2317.4 202960.0 Colocasia esculenta 30.1 11872.0 108.6 19530.8 43830.0 Cyrtosperma merkusii nd nd 18.3 nd 43830.0 Dioscorea alata 39.5 16683.0 0.7 21595.1 nd Dioscorea nummularia 39.5 16683.0 0.7 8867.0 nd Musa sp.1 30.1 9425.0 20.3 1404.4 183.0 Musa sp.2 30.1 9425.0 20.3 1404.4 183.0 Supplementary Amorphophallus paeoniifolius nd 854.1 nd nd nd Dioscorea bulbifera 39.5 854.1 0.7 201.0 nd Inocarpus fagifer nd nd 0.7 57.7 nd Piper methysticum nd nd nd 2527.5 nd Spondias dulcis nd 60.0 nd 5.0 nd Zingiber zerumbet 30.1 nd nd nd nd Average 31.1 6925.0 26.3 5275.0 48528.2

Table 9.4 Labour inputs for species identified at Talasiu modelled using comparative systems

Labour Inputs (hours or instances of activity) Example system Pooled standard deviation Pooled standard error Difference Confidence Mean difference Gadio Enga (primary crops vs supplementary) 9.3 7.6 0.6 <50% 4.9 Bellona Island (primary crops vs supplementary) 5424.6 4429.2 2.4 95% 9050.8 Anuta Atoll (primary crops vs supplementary) 13.4 10.6 0.9 60% 9.8 Tongatapu (primary crops vs supplementary) 7192.8 4650.4 1.6 80% 7192.8 Ontong Java (primary crops vs supplementary) 87899.2 0.0 0.0 0 0.0

Table 9.5 Statistical comparison of labour inputs for groupings at Talasiu in terms of mean difference modelled using comparative systems

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Modelling Leka (J17) The Formative Period (1500-1000 BP) led to the later development of a complex social hierarchy in Tongan prehistory but the archaeological record is characterised currently by a general lack of evidence for any cultural, political or technological change. With regard to changes in subsistence strategies, few sites have been dated to this period, and fewer still have been excavated to collect data on both faunal and floral components of diet. Occupation at Leka (J17) is an important site to assess both these components in terms of the expansion of territorial resource use, due to the decline in available near-shore marine species through overuse and environmental change. The faunal remains from Leka have been analysed by Grono (2012) and confirm that the hydro-isostatic drawdown of sea level after the mid-Holocene reduced the availability of many larger shellfish species, prompting a gradual focus inland that is also reflected within settlement patterns during this time (Poulsen 1967; Spenneman 1986, 1989). The debate surrounding post-Lapita plant production systems, which has focussed on factors such as mental templates, continued interaction and migration, environmental thresholds and change, culturally-created thresholds, and demographic change, has already been outlined. It is important to discuss early production at Leka, and how this compares with expectations based on this contextual background.

The list of identified species in the micro- and macrobotanical remains from Leka again includes a number of arboreal, root and tuber crops, as well as supplementary species that would have been semi-cultivated or gathered from native or naturalised plants. As was the case with species identified in deposits at Talasiu, these are mostly prehistorically introduced plants apart from Cocos nucifera, and all but the White potato (Solanum tuberosum) are considered to have been introduced prior to European contact. This species has been labelled as a contaminant in samples from Leka that probably occurred during laboratory processing but this will be discussed later in this chapter. Piper methysticum is thought to be a late prehistoric introduction, and so is also possibly a contaminant within these samples, but as was the case for Talasiu, this species will be incorporated as a supplementary species as it may be an early introduction. Because so little is known about the Formative Period in Tongan prehistory, none of these species have previously been identified within palaeoenvironmental or archaeological deposits from this time, and therefore these contribute greatly to knowledge upon subsistence and the role plants played in the evolution of social hierarchy and political economy.

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Family Species Anacardiaceae Spondias dulcis

Amorphophallus paeoniifolius Araceae Colocasia esculenta Cyrtosperma merkusii

Arecaceae Cocos nucifera Dioscorea alata Dioscorea bulbifera Dioscoreaceae Dioscorea esculenta Dioscorea nummularia Fabaceae Inocarpus fagifer Lecythidaceae Barringtonia asiatica Moraceae Artocarpus altilis Musaceae Musa spp. Piperaceae Piper methysticum Solanaceae Solanum tuberosum Zingiberaceae Curcuma longa

Table 9.6 Identified families and species within archaeobotanical remains from Leka (J17)

The combined list of species from Leka was modelled as a complete plant production system and compared with the range of modern production systems The same criteria used to characterise modern systems were applied to the data from Leka, and used to gauge the extent of species and overall system diversity and efficiency in terms of nutrition, yield and labour inputs. There are limitations to this, as some plant data is not available for all modern systems, but when these species are treated as a system, patterning emerges that at least eliminates some modern systems and pinpoints others that are more likely similar to that at Leka. This system has moderate species diversity in all primary and supplementary species (16 species in total), but insignificant nutritional diversity between these groupings (<50% confidence). The labour and yield diversity of species in the Leka system cannot be known. However, when these species are compared using the figures from all of the example systems, a range of possibilities can be gauged. The average yield ratios for species within the Leka system can vary from low- to-high diversity (see Table 9.7), when modelled using figures from all of the comparative systems. Average labour input figures for each grouping varied markedly between these example systems (see Table 9.8), but most were not able to be shown to be statistically different using Student’s t test and associated two-sided distribution. Labour diversity between the groupings was only significant when modelled using data from the Bellona system (2.64 standard deviations and 95% confidence), but was followed closely by Anuta (0.86 standard deviations and 90% confidence). From these descriptions the Leka system can be characterised as:

 Leka— moderate species diversity, insignificant nutritional diversity between groupings, significant labour diversity, low-to-high yield ratio diversity.

245

The description of the Leka system, if the list of species identified archaeobotanically within test units at this site are considered to represent a complete system, most closely resembles the Anutan system. The data recorded and compared for the Anutan system suggested that the utilisation of plants on this small high island involved a moderate number of species, but when these were grouped into primary or supplementary species according to the nature of their use and production there was no significant nutritional diversity between these groups. However, there was significant diversity within labour inputs for each of these groups, which differed by 3.17 standard deviations with 100% confidence. Overall yield ratio diversity was low between species within the Anutan system. If this model is correct, then the overall efficiency of the Leka system was 1268kcal/hr, which is the least efficient of all of the simulated systems for Leka, and is significantly lower than Talasiu. This exercise in modelling the archaeological production system at Leka does not meet the original expectations, which suggested that this system would most closely resemble the Bellona system based on current hypotheses for island colonisation and the role of agriculture in the development of social hierarchy. If this model is correct, then the Leka system was probably more labour intensive than originally presumed, and focussed on the cultivation of a narrow range of primary crops, with little labour invested in the exploitation of supplementary species. The implications of these findings will be discussed later in further detail.

Species Gadio Enga Bellona Is Anuta Tongatapu Ontong Java Amorphophallus paeoniifolius 0.0 2.9 0.0 0.0 0.0 Artocarpus altilis 1.0 41.7 0.1 0.0 0.0 Cocos nucifera 0.0 30.0 0.0 0.0 0.5 Colocasia esculenta 66.1 5.5 0.0 8.9 0.9 Curcuma longa 0.0 0.0 0.0 0.0 0.0 Cyrtosperma merkusii 0.0 0.0 0.5 0.0 1.6 Dioscorea alata 0.3 12.7 0.0 12.1 0.0 Dioscorea bulbifera 0.3 2.9 0.0 12.1 0.0 Dioscorea esculenta 0.3 12.7 0.0 12.1 0.0 Dioscorea nummularia 0.3 12.7 0.0 12.1 0.0 Inocarpus fagifer 0.0 0.0 0.0 0.0 0.0 Musa sp.1 5.7 21.4 2.5 25.1 9.6 Musa sp.2 5.7 21.4 2.5 25.1 9.6 Piper methysticum 0.0 0.0 0.0 0.0 0.0 Solanum tuberosum 0.0 0.0 0.0 0.0 0.0 Spondias dulcis 0.0 83.3 0.0 0.0 0.0 Average 5.0 15.5 0.3 6.7 1.4

Table 9.7 Yield ratios for species identified at Leka modelled using comparative systems

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Gadio Enga Bellona Is Anuta Tongatapu Ontong Java Grouping Species (instances of activity) (hours/yr) (hrs/37 days) (hrs/yr) (hrs/yr) Primary crops Artocarpus altilis 10.3 60.0 62.6 115.2 183.0 Cocos nucifera nd 3333.3 30.3 2317.4 202960.0 Colocasia esculenta 30.1 11872.0 108.6 19530.8 43830.0 Cyrtosperma merkusii nd nd 18.3 nd 43830.0 Dioscorea alata 39.5 16683.0 0.7 21595.1 nd Dioscorea esculenta 39.5 16683.0 0.7 8867.0 nd Dioscorea nummularia 39.5 16683.0 0.7 8867.0 nd Musa sp.1 30.1 9425.0 20.3 1404.4 183.0 Musa sp.2 30.1 9425.0 20.3 1404.4 183.0 Supplementary Amorphophallus paeoniifolius nd 854.1 nd nd nd Curcuma longa nd nd nd nd 9200.0 Dioscorea bulbifera 39.5 854.1 0.7 201.0 nd Inocarpus fagifer nd nd 0.7 57.7 nd Piper methysticum nd nd nd 2527.5 nd Spondias dulcis nd 60.0 nd 5.0 nd Zingiber zerumbet 30.1 nd nd nd nd Contamination Solanum tuberosum nd nd nd nd nd Average 32.0 7812.0 24.0 5574.4 42909.9

Table 9.8 Labour inputs for species identified at Leka modelled using comparative systems

Labour Inputs (hours or instances of activity) Example system Pooled standard deviation Pooled standard error Difference Confidence Mean difference Gadio Enga (primary crops vs supplementary) 9.3 7.4 0.5 <50% 3.5 Bellona Island (primary crops vs supplementary) 5565.9 3768.1 2.6 95% 9931.1 Anuta Atoll (primary crops vs supplementary) 13.0 10.2 0.9 50% 8.7 Tongatapu (primary crops vs supplementary) 7044.6 4313.9 1.7 80% 7314.9 Ontong Java (primary crops vs supplementary) 78619.5 0.0 0.0 0 48528.2

Table 9.9 Statistical comparison of labour inputs for groupings at Leka in terms of mean difference modelled using comparative systems

Modelling Heketa (TO-Nt-2) The cultural deposits at Heketa have radiocarbon ages which place occupation at this site to the Formative Period, and transitioning to the early stages of the Classic Tu’i Tonga chiefdom initially based at Heketa before moving to south to Lapaha after the 14th century AD. Two distinct cultural deposits were dated using AMS. Charred coconut endocarp from Layer 3 dated to around 600 cal BP, while endocarp from Layers 4-5 produced a calibrated range of 800-600 cal BP. There is some overlap between these ranges, and so it is possible that these represent either two discreet systems or events, or a continuity of site occupation and associated plant production. When the ranges of identified species within these deposits are compared, there are differences in the number and type of species present. Layer 3 has ten species including six primary and four supplementary species. Layers 4-5 have seven species, which includes five primary and only two supplementary species. Due to the possible overlap in dates and low overall preservation of microbotanical remains within these deposits from Heketa TP4 that may have biased the distribution of these identifications, it was decided to treat the list of species identified within TP4 at Heketa as a single system for modelling. The data provides an interesting insight into the type of production system that may have enabled the creation of 247 surplus for tributes that supported a centralised political system and an archaic state evolving at this time.

A number of arboreal, root and tuber species composed this archaeological production system at Heketa. As was the case for Talasiu and Leka, these species represent a mix of both primary crops and supplementary species. However, the number of species identified within micro- and macrobotanical remains is much smaller compared to the two other sites, with only seven primary crops and four supplementary species. This could suggest a preservation bias. These taxa are primarily prehistoric introductions, aside from Cocos nucifera. Ipomoea batatas is either a prehistoric or historic import to Tonga, but is treated as part of the primary crops within the assemblage from Heketa due to the late prehistoric date ranges for these deposits. This decision was made to at least attempt to model a system that includes I. batatas as this is a dryland crop that currently grows well in Tonga, and if it was introduced earlier than expected, would have played an important role in plant production and overall system efficiency. This will be discussed in greater detail later in this chapter.

Family Species Anacardiaceae Spondias dulcis Alocasia macrorrhiza Amorphophallus paeoniifolius Araceae Colocasia esculenta Cyrtosperma merkusii Arecaceae Cocos nucifera Convolvulaceae Ipomoea batatas Fabaceae Inocarpus fagifer Moraceae Artocarpus altilis Musaceae Musa spp. Piperaceae Piper methysticum

Table 9.10 Identified families and species within archaeobotanical remains from Heketa (TO-Nt-2)

Species diversity at Heketa is very low, with only 11 species in total, again possibly indicating preservation bias. When the taxa are grouped into primary and supplementary species, there is no significant nutritional diversity between these two groups. The averages of these groups differ by 1.56 standard deviations, but based on the number of samples being compared this figure equates to an 80% probability that these samples are different, which is not high enough to reject the null hypothesis that these groups share similar nutritional values. The average yield ratios indicate low-to-high diversity within these species (see Table 9.11), based on the possible range provided by the example systems. Finally, a statistical comparison of labour invested into the primary and supplementary species using Student’s t test, based on the range of data from modern systems, indicates that there is insignificant labour diversity between these groups (see Table 9.12 and Table 9.13). The highest probability of difference was seen within the data from the Bellona system, which differed by 1.3 standard deviations and 70%

248 confidence in mean difference, but again this was not enough to reject the null hypothesis. From these criteria, the archaeological system represented by archaeobotanical data from Heketa can be characterised as:

 Heketa— low species diversity, insignificant nutritional diversity between groupings, insignificant labour diversity between groupings, low-to-high yield ratio diversity.

It was expected that an archaeological plant production system dated to around 800-600 BP in Tonga would most closely resemble the Anutan system, based on current ideas upon the links between intensive systems and the development of social hierarchy. The modelled characterisation of the system at the Heketa site would suggest that this system probably was more like that recorded on Ontong Java by Bayliss-Smith (1973, 1977). If this is true, then the overall efficiency of the Heketa system in terms of outputs to inputs would be higher than if this system had been more similar to Anuta. The result of this is a modelled overall nutritional efficiency of 7846kcal/hr (Ontong Java), rather than 881kcal/hr (Anuta). This characterisation suggests that the Heketa system could have been less labour intensive than originally thought. It is also possible that increased trade networks during this time had enabled the economy to incorporate the export of some higher yielding but lower labour investment crops in exchange for the import of other subsistence items, as seen in the Ontong Java economy.

Table 9.11 Yield ratios for species identified at Heketa modelled using comparative example systems

Gadio Enga Bellona Is Anuta Tongatapu Ontong Java Grouping Species (instances of activity) (hours/yr) (hrs/37 days) (hrs/yr) (hrs/yr) Primary crops Alocasia macrorrhiza nd 11872.0 0.7 296824.2 183.0 Artocarpus altilis 10.3 60.0 62.6 115.2 183.0 Cocos nucifera nd 3333.3 30.3 2317.4 202960.0 Colocasia esculenta 30.1 11872.0 108.6 19530.8 43830.0 Cyrtosperma merkusii nd nd 18.3 nd 43830.0 Ipomoea batatas 30.1 33000.0 0.7 6394.5 183.0 Musa sp.2 30.1 9425.0 20.3 1404.4 183.0 Supplementary Amorphophallus paeoniifolius nd 854.1 nd nd nd Inocarpus fagifer nd nd 0.7 57.7 nd Piper methysticum nd nd nd 2527.5 nd Spondias dulcis nd 60.0 nd 5.0 nd Average 25.1 8809.6 30.9 36575.2 41621.7

Table 9.12 Labour inputs for species identified at Heketa modelled using comparative example systems 249

Labour Inputs (hours or instances of activity) Example system Pooled standard deviation Pooled standard error Difference Confidence Mean difference Gadio Enga (primary crops vs supplementary) 12.1 0.0 0.0 0% 0.0 Bellona Island (primary crops vs supplementary) 10520.1 8589.6 1.3 70% 11136.7 Anuta Atoll (primary crops vs supplementary) 15.2 16.2 0.7 50% 10.8 Tongatapu (primary crops vs supplementary) 100536.9 71090.3 0.8 50% 53567.7 Ontong Java (primary crops vs supplementary) 878.899.24 0.0 0.0 0 0.0

Table 9.13 Statistical comparison of labour inputs for groupings at Heketa in terms of mean difference modelled using comparative systems

Comparison of expected and modelled outcomes There are some significant discrepancies between the expected and modelled characterisations of the archaeological production systems. Based on previous data and hypotheses, it was expected that the late Lapita site of Talasiu would resemble the ‘broad-spectrum’ subsistence system of the Gadio Enga, the Formative Period site at Leka would have been similar to the mixed shifting cultivation of the Bellona system, and the late Formative and early Classic Tu’i Tonga chiefdom site of Heketa would resemble the more intensive Anutan system. However, modelling these archaeological systems based on the criteria for productivity and diversity used to describe modern systems indicated similarities to different systems within this range. The modelling exercise suggested that Talasiu could have instead resembled the Tongatapu 2001 system, while Leka was modelled to have been more labour intensive and less efficient than previously assumed and so closer to the Anutan system. Interestingly, Heketa was suggested to be closer in similarity to the production system of Ontong Java, which features land and labour intensive subsistence and cash-cropping techniques with little investment in gathering of semi- cultivated, naturalised or native species.

Unfortunately, there was no descriptive context provided in the Tongan Agricultural Census (Ministry of Agriculture and Forestry 2001). Therefore, it may be useful to consider the context of the Bellonan system, based on already established similarities between the environmental context, shifting cultivation practices and productivity of these two systems. The Bellona system was based on ‘manipulated and natural ecosystems’ (Christiansen 1975:29). Gardening was conducted alongside gathering, collecting and fishing. The term ‘gardening’ was used by Christiansen for these reasons: “...tilling is not practised, plants are individually placed in individually treated sites, the areas planted are usually small, and generally the plants used in these areas are annuals, vegetatively propagated (corms, tubers, bulbils, cuttings) and are thus not sown, but laid/planted.” (Christiansen 1975:30) Local gardening was described as a rejuvenation of a natural juvenile ecosystem after initial clearing, with high biomass production and relatively low species diversity. Soil fertility was maintained through long fallow periods enabling wild plant regeneration and nutrient cycling. This characterises shifting cultivation and thus the Bellonan system is described as such by Christiansen (1975:31). 250

The exploitation of a number of niches in these manipulated and natural ecosystems was enabled through the application of simple technology that involves low and ‘rational’ inputs of labour to achieve subsistence sufficient only for local needs (1975:30). Christiansen ascribes lower efficiency, but greater stability to the utilisation of natural potential on the island. Analysis of output to input ratios within this study indicated that this was not always the case. For example, Spondias dulcis or Otaheite apple, which is a semi-cultivated species, was shown to have the highest nutritional return ratios of all of the species in the Bellonan system, due to very low labour investment for a high yield. These differences in views on efficiency could be due to the types of outputs measured, seasonal variation or the nature of data included in the original study which also made use of fishing records.

The techniques described for cultivation through shifting cultivation, combined with the exploitation of natural resources on this raised limestone island, makes sense when compared with the Talasiu data. Although it was expected that the balance of labour invested in cultivated to supplementary-type species exploited would be more similar to the Gadio Enga ‘mixed’ subsistence, by 2650 BP the island may have been settled for several hundred years. It is possible that this later stage of island colonisation was already characterised by greater emphasis on multi-cropping in vertically stratified gardens of crop storeys starting with vegeculture interspersed with tree crops such as bananas and breadfruit (Addison 2006) using shifting cultivation.

A more surprising outcome of modelling was the suggestion that the archaeological production system represented by botanical remains extracted from Leka was most similar to the Anutan system. The use and cultivation of plants on Anuta was described by Yen (1973b) as “...one of the most intensive extant in the Pacific, despite the shortage of land and water, which could have conferred the potentiality for the well-known forms of intensive agricultural production, irrigation farming of taro.” Yen (1973b:139) clarifies that this system is a mixed agricultural form of both high and low labour requirements which together indicates a highly intensive system. Technologically, agriculture was seen as intensive due to the construction of structures such as dry terraces or permanent fields, also in terms of land use within agronomy through crop rotation, mulching, and finally labour expenditure. The intensity of this system was such that Yen (1973b:148) also argues that the attainment of carrying capacity would span a shorter period on Anuta than in many other island settings, and in fact the island was close to this point when the original study was conducted. Forms of storage were utilised for many of the root, tuber and tree crops that are high in starch content. This involved the fermentation of processed taro, manioc, breadfruit, banana, Cyrtosperma taro, and Burckella fruits, and sealing within lined pits for periods of time that are crop-specific. Yen describes storage as a possible means of dealing with population increase, and indicates the emergence of more formalised agricultural organisation in Anutan prehistory (1973b:147). Anuta was completely self- 251 sufficient at the time that this system was recorded, without any real cash objectives in agriculture such as those seen elsewhere on the production and export of copra. This presented Yen (1973b) with a unique opportunity to study traditional subsistence without any modern industrial influences, and is one of only two systems analysed within this study that had no cash-cropping.

Several key concepts within Yen’s description of the Anutan system offer possible explanations for the similarities between this and the archaeological system at Leka. At first glance, it is surprising that agriculture in Tonga may have intensified to the point of comparatively low system efficiency in terms of nutritional returns already by 1300-1000 cal BP. However, when this is considered in terms of the modelled carrying capacity of Tongatapu in terms of agricultural acreage required to support an increasing population, argued to have been reached as early as 800 BP (Kirch 1994) or 1000 BP (Green 1973), some contextual similarities emerge. The described self-sufficiency of the Anutan system may have also characterised agricultural production on Tongatapu during dates represented within deposits at Leka, as the extensive networks of trade and interaction associated with the maritime chiefdom were established at this time but later intensified (Clark et al. 2014). It is therefore possible that Tongatapu had been nearing the attainment of carrying capacity at this time, and therefore agricultural production was reducing in system nutritional efficiency during the Formative Period.

This assumption ties into Kirch’s (1994) argument that the stimulus for warfare and conquest was often the need to expand agricultural production in dryland systems such as that seen on Tongatapu and in the Hawaiian archipelago. Expansion of political control acquired greater land under one centralised polity, and relieved pressure on existing land currently under production. The development of political economies, especially within primary and secondary states, is therefore generally seen as directly related to the emergence of intensive and expansive agricultural systems. Kirch (1994:27-244) drew on the ethnographic as well as archaeological record of the two chiefdoms of Sigave and Alo-Alofi from Futuna, which indicated that agricultural practices in wet and dry environments responded to identical underlying pressures of population increase and of the societal demands for surplus to fuel the competitive exchange/feasting cycle. Whereas Sigave (wet) could utilise opportunities for pondfield irrigation of taro, the opposing chiefdom of Alo-Alofi (dry) was forced to adopt more labor- intensive short-fallow shifting cultivation, closer to Boserup’s model of intensification (1994:244). In an extension upon this model, McCoy and Graves (2012) suggest that the limited potential for surplus production from small valley irrigated fields on the island of Hawaii could place these systems in the same category as rain-fed dryland farming, influencing decisions to engage in expansionist warfare by chiefs seeking to expand their power base. The Hawaiian data is further supported by archaeological and ethnographic research conducted within the 252

Austral archipelago (Bollt 2012), where chronologies of inter- and intra-island warfare were stimulated by the limited potential for irrigation of taro due to island size and environmental constraints.

It is possible that greater investment of labour into primary crops may have reduced overall system efficiency in the past, but energy storage through in-ground surplus accumulation (Bayliss-Smith and Hviding 2014) may have instead increased the efficiency of particular taxa in terms of social returns. These social returns would have both stimulated and been created by the increasing socio-political complexity of the Tongan cultural system. Elites mobilised subsistence goods from commoners by asserting ownership over agricultural land, thus enabling the supply of food to specialists such as warriors and priests who validate the authority of those elites. This Formative Period has been pinpointed as a critical stage in Tongan prehistory that likely lead to the development of the classic Tu’i Tonga chiefdom after 1000 BP, or possibly later. The modelled changes in agricultural production and associated environmental constraints described here could go some way towards explaining why these socio-political developments occurred later.

This hypothesis is further supported by the modelled similarities between the Ontong Java and the archaeological plant production system at Heketa from 800-600 cal BP. It was expected that this archaeological system would most likely resemble the Anutan system due to increasing system intensity over time as a result of the need for surplus production facilitating socio-political developments. However, modelling suggested that in fact the earlier system at Leka was more similar to the Anutan system, and that Heketa most likely resembled the slightly more nutritionally efficient Ontong Java system. Considering this and earlier discussion upon the limits of carrying capacity on Tongatapu, it is possible that by 800-600 cal BP archaeological production had increased in efficiency through the introduction of export items from trade within the increasing network of islands under the control or influence of the maritime chiefdom. Primary crops that were more nutritionally efficient in terms of labour investment could have been incorporated into the diet on Tongatapu, not necessarily through production but through importation and redistribution. This was documented ethnographically as part of the festival and tribute system, where yam production in particular was monitored by chiefly-appointed petty officers and brought to Tongatapu from as far afield as Vavau as part of the first fruits or ‘inasi festival (Cook in Beaglehole 1969; Gifford 1929; Mariner in Martin 1991). A crop of yams was planted around one month before the regular crop, and was harvested early in time to be given as tribute for the festival (Mariner in Martin 1991). Other festivals were also conducted throughout the year to ensure the productive success of cultivation, with gifts of yams, coconuts, kava root, fish and arrowroot given to the gods, the chiefs and their households (Mariner in Martin 1991).

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If this hypothesis is accepted, there are three explanations as to why the system at Heketa may have been similar to the Ontong Java production system, and appears to have increased in overall nutritional efficiency since the Formative Period. The first possibility is that the low preservation of microbotanical remains at Heketa led to the exclusion of some species from the system modelling exercise. The second option is that the incorporation of tributes from other islands, as well as traded goods, allowed renewed investment of labour into more nutritionally efficient but lower yielding species. Tied to this is a third explanation— that the archaeobotanical record at Heketa may not in fact reflect only those species that were produced or utilised from Tongatapu, but also these other islands and so appear greater in overall nutritional efficiency. Each of these will be explored here to further understand and explain the modelling of this archaeological system.

The comparatively low quantities of starch and also the number of species present within the archaeobotanical remains at Heketa suggest that the microbotanical preservation at this site was not good. Most surprising was the absence of any starch identified as belonging to the Dioscorea or yam family. Clearly these were heavily relied upon at the point of European contact, and evidence from Talasiu and Leka indicate that these crops had been cultivated in the past. This suggests that there are certainly some species missing from the production system at Heketa. As yams are not as nutritionally efficient as many other staple crops such as bananas (Musa spp.) or sweet potato (Ipomoea batatas), but are generally more efficient than the aroids, it is possible that the actual production system at Heketa may have been less efficient than that from Ontong Java. Interestingly, yams were not cultivated within the Ontong Java system, which instead focussed primarily on the subsistence production of aroids such as Colocasia esculenta and Cyrtosperma merkusii, sweet potato (Ipomoea batatas) and the cash-cropping of Cocos nucifera. This omission may have biased the model towards characterising the Heketa system as most similar to that from Ontong Java but also may point to the increasingly important role of sweet potato in the Tongan economy.

Overall the two key differences between the Anutan and the Ontong Java system are that there is more labour invested in supplementary species and greater diversity within yield ratio in the latter example system. As has been demonstrated within this study, the investment of some labour into non- or semi-cultivated species often results in high nutritional returns due to high nutritional rankings, especially arboreal species. By specialising in the cultivation of primary crops such as yams and aroids which have moderate yield ratios but low overall nutrition, the Anutan's decrease the overall nutritional efficiency of the system. The type of ‘technoenvironmental efficiency’ which has been measured within this study applies specifically to the Ontong Javan, Bellonan and Tongan ethnographic systems due to the inclusion of de facto outputs. These are the surplus outputs or yields that are created to facilitate the exportation of goods for cash-cropping, but do not include the cash or traded goods that 254 come back into the system. It is therefore possible to measure the role that the creation of surplus would have played, but difficult to accurately assess how the incorporation of any tributes that fed back into the system would have benefitted the efficiency of the production system at Heketa.

It seems likely that the introduction of alternative sources of nutrition without any primary labour inputs would have increased overall efficiency, but this cannot be modelled here without knowing the exact labour investment data. In any case, it is unlikely that goods were only moving in one direction— into Tongatapu—after the Fiji-Western Polynesian interaction sphere was established. The incorporation of tributes into diet and the economy on Tongatapu may have enabled the renewed investment of labour into more nutritionally efficient supplementary species and therefore may explain the increase in system efficiency after the Formative Period (Leka). We cannot know for sure whether species present at Heketa were grown nearby but, as explored here, this may explain the similarities to Ontong Java along with the changes in decision-making in production resulting from the tribute system.

Specialisation and system efficiency Patterning between modelled system efficiency and ratios of numbers of primary to supplementary systems indicates a link between these two variables. Modelled system nutritional efficiency was based on figures derived from the characterisation of past systems and similarities to modern production systems (see Figure 8.36). Talasiu most closely resembled the system on Tongatapu in 2001, while Leka resembled Anuta and Heketa probably resembled the Ontong Java system (see Table 9.14). The modelled efficiency of these systems therefore decreases between Talasiu (2750–2650 cal BP) and Leka (1300-1000 cal BP), before increasing slightly again at Heketa (800-600 cal BP). The numbers of primary and supplementary species within these systems vary over time. The highest ratio of primary to supplementary species was observed at Leka (2:1), while the lowest was at Talasiu (1.3:1). Considering this, a link can be drawn between specialisation in primary crops, seen through this high ratio at Leka, and decreased system nutritional efficiency. Similarly, diversification through exploitation of a number of supplementary taxa alongside primary crops, can be linked to increased system efficiency at Talasiu, with Heketa falling somewhere between these two systems.

Archaeological Ratio of primary to Modelled system Nutritional efficiency system supplementary species (kcal/hr) Talasiu (2750-2650 cal BP) 1.3:1 Tongatapu 123996 Leka (1300-1000 cal BP) 2:1 Anuta 1268 Heketa (800-600 cal BP) 1.4:1 Ontong Java 7846

Table 9.14 Comparison of modelled system efficiency with ratios of primary to supplementary species

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Contamination at Leka and Heketa The issue of contamination from white potato (Solanum tuberosum) and also possibly sweet potato within archaeological deposits at Leka and Heketa needs to be addressed. Contamination is most likely to have occurred during field collection, or laboratory processing for starch extraction. Modern starch contamination during collection and handling of samples in the field has been discussed within a number of studies. These have indicated that modern starch can enter archaeological soil samples during sampling from aeolian (wind) sources (Laurence (2013; Laurence et al. 2011; Loy and Barton 2006; Messner 2011), although this is unlikely in Tonga as starch tends to only becomes airborne when dried, or from handling of samples (Barton et al. 1998:1233; Hart 2011) Through experimentation with laboratory processing and also assessment of current literature, it was discovered that some laboratory grade chemicals and equipment contained modern native starch. Previous studies had recorded high numbers of starch from wheat (Triticum sp.), white potato (Solanum tuberosum), and maize (Zea mays) on powder-free gloves, pipette tips, paper towels, and within Calgon and Sodium polytungstate commonly used for heavy liquid separation (Campbell et al. 1984; Crowther et al. 2014; Laurence 2013; Laurence et al. 2011; Makela et al. 1997; Newsom and Shaw 1997; Wilson and Garach 1981). This was confirmed by testing lab materials used in this study, with Calgon providing the greatest source of contamination. Airborne starch was also found in laboratory spaces in low numbers in native and modified condition (Crowther et al. 2014; Laurence et al. 2011; Newsom and Shaw 1997).

Appropriate protocols to limit starch contamination have been suggested by previous studies, and were built on here. These involved using starch-free gloves and zip lock bags both in the field and in the laboratory, cleaning the face of the baulk before sampling in the field, cleaning sampling equipment or using individual sampling equipment for each stratigraphic unit, sterilising all laboratory equipment and workspaces, and filtering 5% Calgon solutions to remove any modern starch before use. Unfortunately, these techniques can only limit contamination and it is always possible that some modern starch has entered archaeological samples. However, the common types of contaminants, namely wheat, maize and white potato, would be easily identified within samples from Tongatapu as these are unlikely to have been grown at any time on the island. The presence of the white potato or Solanum tuberosum at Leka is most likely explained as a contaminant from pre- or post-excavation handling or processing.

The historical integrity of samples is therefore presumed to be intact and all other species present are assumed to be of prehistoric age. This is supported by the co-occurrence of parenchyma at Talasiu identified as C. esculenta, A. altilis fruit, D. alata, D. nummularia, Musa spp., and Zingiber spp. It is also assumed here that the presence of sweet potato at Heketa is not a modern contaminant as the dating of other archaeobotanical remains of this species within 256 deposits from Eastern and Central Polynesia coincide with the later deposits from TP4. There are also numerous instances of early historic and ethnographic documentation of extensive sweet potato cultivation on Tongatapu. This evidence will be discussed in detail within the next section of this chapter, dealing with the timing of crop introductions into Tonga.

Linking archaeobotanical data to island colonisation and social complexity The archaeobotanical identifications from Talasiu (TO-Mu-2), Leka (J17), and Heketa (TO-Nt- 2) were discussed in this section, in terms of agricultural production systems at various points in Tongan prehistory, and the broader context of these species-level identifications. Modelling archaeological production systems based on characteristics defined for modern systems in the west Pacific, such as species diversity and nutritional efficiency in terms of outputs to labour inputs, enabled a basic analysis of crop production in the past. Some variables were unknown and so the full possible range for each species, based on the figures from modern systems, was assumed in each archaeological system in terms of yield and labour inputs. Species in each system were defined as primary crops or supplementary based on ethnographic and agricultural studies from Tonga, in order to assess the statistical differences between these two groups in terms of yield, nutritional efficiency and labour inputs.

Comparison of modern systems revealed that overall system efficiency was often tied to the inclusion of a number of supplementary species within subsistence systems, rather than a heavy reliance on primary crops. Specialisation in primary crops can therefore be linked to decreased overall system nutritional efficiency. Supplementary species, especially arboreal, tended to have higher nutritional value than most primary crops, but had lower yield ratios in terms of productivity from labour investment. Some balance between the two groups resulted in higher system nutritional efficiency, such as that seen in the Bellona system in the Solomon Islands (Christiansen 1975). Less efficient systems such as that observed by Yen (1973b) on Anuta invested more labour into a narrow range of primary crops, resulting in lower ratios of nutritional returns to labour inputs than that recorded in other systems. This trend was reinforced through the links between ratios of primary to supplementary species, and modelled system efficiency within the archaeological systems. This patterning was critical towards linking decreased system efficiency to the formation of centralised government on Tongatapu.

The results of this study suggest that plant production strategies after initial settlement on Tongatapu by the eastern Lapita Cultural Complex, under a decentralised social hierarchy, may have involved a greater diversity of species than originally presumed. Many of the core traditional cultigens were present within archaeological deposits at Talasiu from 2750-2650 cal BP, including a number of root crops from the aroids (Araceae), and yams (Dioscorea spp), as well as arboreal crops such as bananas (Musaceae) and breadfruit (Artocarpus altilis).

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Alongside these were a number of supplementary crops such as kava (Piper methysticum), ginger (Zingiberaceae), and arboreal fruits including the Otaheite apple (Spondias dulcis) and the Tahitian chestnut (Inocarpus fagifer). This range of both primary and supplementary species most likely resulted in a relatively efficient system in terms of nutritional returns through diversification of labour investment, and would have been utilised alongside hunting and gathering of marine resources. The hydro-isostatic drawdown affecting sea levels within the Fanga’Uta Lagoon during the mid-Holocene (Dickinson 2001, 2007) appear to have gradually impacted this harvesting of marine species in terms of lower species diversity and size (Grono 2012). Overall subsistence strategies at this time appear to have turned towards the exploitation of terrestrial resources in the archaeobotanical record at Leka from 1300-1000 cal BP. The number of species present within test units at this site increase slightly from that observed at Talasiu to include a number of highly ranked crops such as Dioscorea esculenta and Curcuma longa in terms of nutritional value but low in terms of yield to labour investment. These species served to increase the diversity of labour investment between primary and supplementary species (suggesting specialisation), and decrease the overall efficiency of the system. Finally, a much smaller list of species was recovered from archaeological deposits under the Heketa (TO- Nt-2) in the north east of Tongatapu. This is thought to be due in part to taphonomic factors affecting microbotanical preservation, but also could reflect subsistence changes resulting from the development of social hierarchy in Tongan. Overall, modelled system efficiency increased slightly from that seen at Leka, owing to less labour diversity within primary and supplementary crops (diversification) and higher overall yield to labour input ratios. It has been suggested that the networks established through the maritime chiefdom would have increased trade and tributes coming into Tongatapu, alleviating pressure on subsistence when carrying capacity was approached.

These plant production systems were clearly engaged in a cyclical system of involvement with the evolution of social hierarchy and associated socio-political development in Tonga over time. They both facilitated and encouraged change through cultural reactions to decreasing system nutritional efficiency, and were impacted by these changes through the introduction of new species or changing species diversity (diversification vs. specialisation). The assessment of individual species introductions into Tonga further expands the context of these changes. Most species were introduced into Tonga within the first few hundred years of settlement, relied on as supplementary or primary cultigens that complemented the exploitation of marine resources. Demographic and environmental factors such as soil geology and climatic limitations are thought to have pushed Tongatapu to reach carrying capacity by as early as 1000 BP, necessitating expansion of political control to alleviate pressure on local subsistence systems. Clearly, there is an overall decrease in system nutritional efficiency and increase in

258 social complexity after 2750-2650 cal BP, linked to increased specialisation in the production of primary crops over supplementary crops (see Figure 9.1).

Figure 9.1 Trend towards decreased system nutritional efficiency and increased social complexity after Lapita occupation at Talasiu (2750-2650 cal BP)

Modelling archaeological production systems and the introduction and utilisation of individual species into Tongatapu was carefully assessed with relation to current debates and evidence from both Tonga and the wider Pacific. Attempts were made to model how and why plant production systems were utilised on Tongatapu, using high resolution techniques to extract archaeobotanical data from three well-dated sites. The data from Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2) was interpreted within a Human Ecological framework to model agricultural systems, and sought to validate or provide alternatives to current hypotheses and expectations. The implementation of further archaeobotanical research will either support or disprove the narrative for the development of agricultural systems in Tongan prehistory presented here, but at least a new baseline has been provided against which new data can be applied.

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Chapter 10 Conclusion Meeting research aims and objectives This thesis had two main objectives. The first of these was to construct a comprehensive comparative collection of both starch grains and vegetative storage parenchyma from economic and supplementary plant species in Tonga. A detailed morphological study was carried out on these micro- and macrobotanical remains to discern how taxa could be differentiated. Morphological analysis also enabled the development of tools (multivariate statistics and identification flowchart keys) to identify samples of unknown origin to taxonomic levels. The second objective of this research was to utilise the comparative collection to identify archaeobotanical remains from three sites on Tongatapu dated from 2750-2650 cal BP (Talasiu), 1300-1000 cal BP (Leka), and 800-600 cal BP (Heketa). Cultural deposits at these sites contain archaeological data from three time periods within Tongan prehistory, ranging from late Lapita through to the early stages of state formation in Tonga.

Several key research questions were asked of the archaeobotanical data from Talasiu (TO-Mu-2), and deposits underneath Langi leka (J17) and Heketa (TO-Nt-2). One primary question examined was whether early colonisers in West Polynesia were dependent on introduced crops, or if human dispersal was fuelled predominantly by the exploitation of natural resources. A further research question sought to establish whether archaeobotanical data can provide new evidence to examine the role of agriculture within the development of the maritime chiefdom in Tonga. After conducting this archaeobotanical research, several important conclusions were drawn about the comparative collection and the role of plants in Tongan prehistory. These are summarised here and will be explored in further detail within this chapter:

 Multivariate statistical analysis and identification flowcharts can enable the discrimination of starch and vegetative storage parenchyma from most Tongan economic and supplementary taxa based on metric and nominal morphological attributes.  A detailed chronology of plant introductions from three sites on Tongatapu indicate that most staple cultigens and some supplementary or famine foods were brought to Tonga within a few hundred years or less of initial Lapita colonisation. Late prehistoric introductions likely included the sweet potato (Ipomoea batatas) by 600 BP, transported from South America via East Polynesia through the extensive trade networks of the developing Tongan state.  Production systems involving the cultivation of these core cultigens alongside a number of semi-cultivated taxa, changed over time in terms of species diversity through diversification of labour or specialisation in primary crops, and overall system efficiency based on modelled nutritional value, yields and labour investment. Modelled 260

production systems here links decreased system efficiency to increased horticultural specialisation. This trend can be linked to a contemporary increase in social complexity through development of centralised government and control of resources that enabled a slight increase of system nutritional efficiency by 800-600 cal BP through minor diversification (either through production on Tongatapu or de-facto outputs). This supports the view that expansion of Tongan control within Fiji-West Polynesia was likely stimulated by extensive dryland agricultural production on Tongatapu, population increase and environmental limitations to production.

One of the primary aims of this thesis research was to develop a comprehensive reference collection of starch and vegetative storage parenchyma from economic plants in Tonga. This was a crucial step towards identifying archaeobotanical remains from the three sites of Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2) on Tongatapu. Around 40 plant species were collected during surveys of plantations in Tonga, Fiji and Palau. Of these, 35 species were able to be sampled and analysed for parenchyma, and 27 contained significant quantities of starch to enable statistical sampling of granule morphology. A number of species had several organs sampled, as more than one organ could be consumed or otherwise utilised according to ethnographic and ethnobotanical records of plant use. To enable accurate identification of these micro- and macrobotanical remains, detailed morphological studies were carried out that assessed a range of both metric and nominal attributes. The use of high resolution imagery with Scanning Electron Microscopy (SEM) enabled greater accuracy of measurement and more detailed description of morphological variables than with standard light microscopy. Morphological studies facilitated differentiation of the species within the reference collection through characterisation of the full range of morphological variation in each sample. There was considerable overlap within the distribution of many of these attributes within both starch and parenchyma in the reference collection, but multivariate statistical comparison of metric variables highlighted attributes where these species varied.

Both starch granules and vegetative storage parenchyma form and compose the structure of plant organs that are most commonly consumed within the traditional subsistence strategies of the Pacific region and represent crucial direct evidence for the cultivation and dietary contribution of Pacific cultigens. It is essential that the morphological characteristics of each species is imaged and described accurately to identify these micro and macrobotanical remains. A combination of light microscopy and SEM were used to image and measure the variables described here, and this data was assessed using a combination of identification flowcharts and multivariate statistics in the form of Discriminant Function Analysis (DFA). All summed data and imagery was entered into a Filemaker database that will be made available online for public access.

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Identification flowcharts are more basic forms of morphological descriptions and were used to distinguish the range of characteristics present within each species or plant organ sampled in the reference collection for parenchyma. Due to the range of characteristics within different tissue types in the samples of parenchyma, it was decided that multivariate statistical analysis of the full range of attributes for each species would not be useful. Tables were created in this study for both starch and parenchyma that summarised a number of key characteristics and will facilitate the identification of unknown archaeological material. From this, two flowcharts were used (with vascular tissues and without) that broke down these attributes of parenchyma within the reference collection. These incorporated both fresh and charred morphological characteristics of the various boundary, ground and vascular tissues. Sometimes it was not possible to distinguish samples beyond family or genus level due to very similar characteristics or a large amount of overlap. Overall, these identification flowchart keys allowed a number of key taxa to be identified with moderate confidence and prefixed with ‘C.f’, as samples both resembled images and matched geographic range of species.

Multivariate statistics enabled a full comparison of all metric and nominal variables within starch granule morphology, and was also utilised to discriminate between the distributions of individual attributes of parenchyma within species in the reference collection. Nominal (discontinuous) variables were converted into binary (presence or absence) variables to facilitate the comparison of these non-metric attributes. A comparison of the full range of morphological variables in starch granules was conducted by dividing the main dataset into two smaller datasets, according to the angle at which granules were viewed and recorded. This created an eccentric (side-on) and a centric (end-on) dataset. Confusion matrices were created that indicated how well each species or plant organ could be distinguished from all other species included within each dataset. These showed the distribution of classifications when linear discrimination is used to assess morphological variance for each species, in essence these highlight the percentage of starch granules that were correctly re-assigned to the species of origin based on the characteristics recorded. Higher percentages of correct re-assignments indicated that a species or organ contains starch that is less similar to other species and can therefore be more easily distinguished morphologically from others within the dataset. These matrices also indicated the species that morphologically overlap and therefore tended be misclassified as one another. The overall results of this analysis produced 67.61% correct re- assignments within the eccentric (side-on) view and 46.78% within the centric (end-on) view.

This technique has both strengths and weaknesses, resulting from inherent statistical assumptions. A number of parameters have an effect on the accuracy rate of DFA classification, including the number of groups, the number of predictor variables, and sample sizes in each group. A primary weakness of the automated classifications generated by DFA is that this technique must provide a group assignment. These assignments are therefore exclusive and so 262 restrict identification of unknown samples only to those groups or species included in the analysis. If archaeological (unknown) starch derives from taxa that are not in the comparative collection, DFA will still attempt to classify granules to taxa or groups that are included. However, a review of the scores of the first two canonical variates for each unknown granule can indicate how close the match is to the range for reference species. DFA is also designed to maximise the differences between groups. This results in an overall tendency to correctly assign individual granules at a higher rate than that expected by chance alone, even if the dataset is comprised of predictor variables (attributes) that in fact bear no real relationship to group membership. Unless each of the outputs for predictor variables are assessed (in this case attributes) the results of DFA can be mistakenly interpreted as representing the successful attribution of individuals to their groups on the basis of meaningful functions, when instead these are the result of the inherent property of the analysis. With this in mind, it is important that the discriminatory values of attributes are assessed before the classification of unknown granules is attempted. This check provides an indication of the probability of failing to distinguish between two plant taxa. Another benefit of this step is that this provides an assessment of all morphological variables and identifies which are most important for discrimination between taxa, and classification. Overall, multivariate statistical analysis created a useful automated classification system for archaeological starch that could be interpreted with low-to-high confidence, and confirmed using visual checking of images from both light microscopy and SEM. Observer bias is not removed from this process but further highlights the importance of using a population rather than a single case approach to identification. Not all granule morphologies are characteristic and the presence or absence of a nondiagnostic form in association with a more generic type could be definitive of a particular species (Torrence et al. 2004), a relationship that only multivariate statistical analysis considers.

This archaeobotanical study is the first to combine two detailed complementary records of micro- and macrobotanical remains from the same organs of economic plant species in the Pacific region. A stringent sampling strategy and consistent recording techniques, particularly in the case of starch, enabled the accurate recording of a number of variables deemed useful within current literature (Crowther 2009; Hather 1991, 1994, 2000; ICSN 2011; Torrence, Wright and Conway 2004; Wilson et al. 2010). This process resulted in the identification of many diagnostic attributes at various taxonomic levels. Further, the incorporation of multivariate statistical analysis highlighted the amount of morphological overlap between these species, and pinpointed those species that could be identified with greater confidence than others within the reference collection. It is intended that one major output of this research will be to make the recorded data and images from both light microscopy and SEM available online, using the Filemaker database that was created during this first component of this PhD project. There are no online databases currently available for the Pacific region, and so this will be a major step

263 towards encouraging more accurate and consistent identifications for preserved starch and parenchymatous material found in palaeoecological and archaeological deposits.

Prior to this thesis research, little was known about the antiquity of crop introductions into Tonga and their use in plant production systems in the past. A small number of palaeoenvironmental studies (Fall 2005, 2010; Fall and Drezner 2011, 2013; Flenley et al. 1999) have painted a picture of vegetation change from pollen and spores within swamp cores, and also explored the timing and mechanisms for plant dispersal. The impact of human arrival in Vava’u, Tonga was demonstrated through an increase in micro-charcoal after 2600 BP suggesting the burning of tropical hardwood, and possible increases in soil erosion from increased clay deposition between 2400 and 800 14C yr BP (Fall 2005, 2010). Lowered numbers of native birds and bats (Steadman 2002) also probably led to the loss of rainforest trees through disrupting seed dispersal mechanisms The introduction of a number of species after 2600 BP, or an increase in pollen concentrations, is thought to be associated with Lapita arrival and later Polynesian occupation (2010). New species introductions around 2600-2500 BP included Casuarina equisetifolia, Colocasia esculenta, Cordyline fruticosa, Ludwigia octovalis, Poaceae spp., and Pometia pinnata (Fall 2005; Flenley et al. 1999). Those species for which pollen increased after this time, suggesting cultivation or expansion of habitat, included Canarium harveyi, Cocos nucifera, and Pandanus tectorius (Fall 2005; Flenley et al. 1999). The first appearance of a number of other species was documented within these studies from 2400- 1100 BP, including Erythrina variegata, Gardenia tannaensis and Stenochlaena palustris (Fall 2005; Flenley et al. 1999).

It is clear that there was a significant gap in terms of data regarding most species that featured within ethnographic recordings of the traditional Tongan economy. Research as part of this project sought to fill this gap through the application of high-resolution archaeobotanical techniques upon well-dated archaeological deposits from throughout Tongan prehistory. Through these methods, a chronology for the introduction and cultivation of many new plant species was developed and can now be added to the existing data. From the site of Talasiu (TO- Mu-2), dated to around 2750-2650 cal BP and therefore associated with late Lapita occupation on Tongatapu, the first recorded evidence for a number of new economic species was documented. These species included the root and tuber crops Amorphophallus paeoniifolius (elephant foot yam or stink lily), Cyrtosperma merkusii (giant swamp taro), Dioscorea alata (common yam), Dioscorea bulbifera (bitter yam), Dioscorea nummularia (spiny yam), Piper methysticum (kava), and several members of the Zingiberaceae family (gingers). Introduced alongside these were a number of arboreal or tree crops including Artocarpus altilis (breadfruit), Inocarpus fagifer (Tahitian chestnut), Musa spp. (banana and plantains), and Spondias dulcis (Otaheite apple). The presence of starch and charred endocarp of both Cocos nucifera (coconut) and Barringtonia asiatica (fish-poison tree) within these deposits indicate that these species 264 were utilised by those populations occupying Talasiu. Only two new species were recorded from deposits at Leka (J17) dated to 1300-1000 cal BP. These included another species of yam, Dioscorea esculenta (lesser yam), and another ginger, Curcuma longa (turmeric). Finally, the introduction of Ipomoea batatas (sweet potato) and Alocasia macrorrhiza (giant taro) can be documented from identified starch granules within deposits dated to 800-600 cal BP under the Heketa (TO-Nt-2).

To understand the development of production systems that featured the cultivation and consumption of these species, and how these linked to social complexity, a Human Ecological approach to modelling archaeological systems was taken. A simple cost-benefit assessment of system efficiency was carried out in terms of work inputs to yield and nutritional outputs, as properties of systems within agricultural ecology (agroecology). Characterisation of these systems were according to criteria roughly based on those suggested by Dornstreich (1977) and included species diversity (number of species utilised), nutritional diversity between groupings (e.g. primary or supplementary species), labour investment diversity between groupings, and yield ratio diversity between all species. System ‘completeness’ was assessed through nutritional comparisons with traditional economic species that were ethnographically recorded at the time of European contact. It was concluded that there was no statistical differences in terms of nutrition between those recorded ethnographically and archaeobotanically, even when these were narrowed to those that produce starch or need to be processed or cooked before consumption, and therefore more likely to be included in midden deposits. The only exception to this is possibly Heketa, where starch quantities were very low compared to the other two sites, and the absence of any yam species at this later stage in Tongan prehistory could suggest that preservation of micro-remains at this site was not good and therefore the record is biased in some way.

These archaeological systems were compared to a small number of published ethnographic examples of systems from locations in the Western Pacific. These included that of the Gadio Enga in the Highlands of Papua New Guinea (Dornstreich 1977), three Solomon outliers including Bellona Island (Christiansen 1975), Ontong Java (Bayliss-Smith 1973, 1977, 1986) and Anuta (Yen 1973b), and Tongatapu (Ministry of Agriculture and Forestry 2001).This exercise suggested that the late-Lapita production system at Talasiu (2750-2650 cal BP) most closely resembled that recorded in the 2001 Tongan Agricultural Census, and therefore had an overall modelled system efficiency of around 124kcal/hr. Plant production during the Formative Period, as represented by identified species at Leka (1300-1000 cal BP) most closely resembled the Anutan production system, and as such had a much lower modelled overall efficiency of 1.3kcal/hr. Finally production at the site of Heketa (800-600 cal BP) during the late Formative/early Classic Tu’i Tonga Chiefdom period could have most closely resembled the Ontong Javan system with an overall system efficiency of 7.8kcal/hr. This indicated that 265 productive and nutritional efficiency decreased between Talasiu and Leka, but increased again between Leka and Heketa, and emphasised the need to assess these systems in historical context.

It is argued here that Lapita colonising populations on Tongatapu likely invested in the production of a more diverse range of traditional core cultigens and supplementary species than originally presumed. The nutritional value of these species and the implied nature of their cultivation using a system of aroid pits (Cyrtosperma) and dryland multi-cropping techniques with elements of arboriculture would have resulted in a relatively efficient system that was used alongside marine-based and terrestrial foraging. Data from Leka suggest that plant utilisation during the Formative Period reflected an increased interest in the exploitation of terrestrial resources. This was due to marine resource over-exploitation and changes in the marine ecosystem within the Fanga’Uta Lagoon resulting from hydro-isostatic drawdown of sea level after the mid-Holocene marine high-stand. Specialisation in primary crops within the production system at this time subsequently increased diversity within labour investment in primary and supplementary crops and decreased overall system efficiency due to the limited nutritional efficiency of new species. Soon after, Tongatapu may have been close to reaching carrying capacity based on modelled population figures and acreages of cultivated land required to feed them (Green 1973; Kirch 1984, 1988). One reaction to this situation would have been the beginning of expansionist strategies that enabled the eventual development of the classic Tu’i Tonga maritime chiefdom or state. The trade networks and control of agricultural resources established by the central polity on Tongatapu after around 750 BP (Clark et al. 2014) created an interaction sphere that may have served to relieve pressure on local production. This could explain the modelled increase in overall system efficiency in the system at Heketa from 800-600 cal BP, owing to less labour diversity within primary and supplementary crops through minor diversification into nutritionally efficient supplementary taxa, and higher overall yield to labour input ratios.

A simple unilinear trajectory towards increasing labour inputs into dryland techniques through fallow reduction and expansion of production, and decreasing overall system efficiency, is not supported by the modelled data from this study. Instead evidence points towards a system of causation and reaction to social, environmental and economic factors that enabled populations to not only survive but also thrive on a raised limestone island such as Tongatapu where a variety of ecological niches for diverse wetland cultivation techniques, such as those observed on high islands, did not exist. The socio-political climate on the island evolved to cope with environmental and climatic limitations on production, through expansion of control and influence.

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Palaeonvironmental and ethnobotanical studies point to the almost complete removal of primary forest on Tongatapu, replaced with plantations in various states of cultivation or bush fallow, or areas of now permanent secondary forest (Ellison 1989; Fall 2005, 2010; Fall and Drezner 2011, 2013. The geology and rich volcanic soils of the island lent themselves to the gradual transformation of the landscape to full scale extensive multi-cropping and shifting dryland production. This process inevitably led to demographic pressure on production, and island carrying capacity being reached earlier than on many other Pacific islands (e.g. Vanuatu, Fiji and New Caledonia). The system could not be further intensified in terms of further fallow reduction or increased cultivation, and so expansion of political control to other islands in the archipelago and the greater Western Pacific region would have been necessary. New interaction spheres and control of resources could have enabled a change in the focus of vegeculture away from some less efficient primary crops towards new species such as Ipomoea batatas (sweet potato) (Allen and Ballard 2001; Ballard 2005; Green 2005; Leach 2005). All producers participated in ethnographically recorded tribute systems (Cook in Beaglehole 1969; Gifford 1929; Mariner in Martin 1991) which created alternative systems of food redistribution. This would have almost certainly nutritionally as well as socially benefitted most elements of society, from the chiefs to the matapules and commoners in different ways. Ultimately, the adaptation of production systems within an evolving social and political climate was a continuing process that did not halt after European contact. The continued ability of those on Tongatapu to adopt new crops such as Xanthosoma sagittifolium (tannia or talo futuna) and increase production of traditional cultigens for export outside Tonga is evidence of a resilient relationship between the people and the land.

Future recommendations

Micro- and macrobotanical techniques A number of techniques were utilised in this study that are currently relatively underemployed in the Pacific. Very few archaeological studies incorporate any archaeobotanical techniques within research strategies, and those that do have tended to be without any strict sampling regimes to ensure consistency and enable replication. Within those studies that have been conducted, there have been numerous studies involving the identification and quantification of pollen, phytoliths and wood charcoal due to high rates of preservation, while very few have targeted starch and even fewer have analysed charred, water-logged or desiccated vegetative storage parenchyma. Background research as part of this study has shown that this bias has been the result of an unfounded belief that these remains do not preserve in tropical climates, and that these remains cannot be confidently identified.

Starch is often analysed only as chance finds as residues on artefacts or soil samples from archaeological features, and parenchyma has similarly been primarily collected as random

267 finds that were often identified based on surface morphology rather than internal cellular structure. These chance finds in the broader Pacific region, and preservation of starch and parenchyma within deposits at Talasiu (TO-Mu-2), Leka (J17) and Heketa (TO-Nt-2), validate the need to incorporate flotation or sieving within excavation techniques in projects that are assessing subsistence and lifeways of people in the past. At the very least, bulk soil samples should be collected to process and analyse later in a laboratory environment where less contamination can occur. This study has demonstrated that these micro- and macrobotanical remains can be extracted from a variety of archaeological deposits, and that these often preserve relatively well in order to facilitate identification using techniques established here. Some consideration does need to be given to sources of contamination within the field and laboratory environment, and taphonomic processes, to ensure the antiquity of identified specimens.

It was foreseen that the development of a comprehensive reference collection would be the best way to understand morphological variation at various taxonomic levels in starch and charred parenchyma, and thus to identify archaeobotanical remains. The analysis of the reference collection built upon the previous work of others (Allen and Ussher 2013; Babot 2001; Barton 2005; Barton et al. 1993,1998; Crowther 2009; Field 2006; Fullagar 2006; Fullagar et al. 1996, 1998; Hather 1991, 1994, 2000; Horrocks 2004;; Horrocks and Barber 2005; Horrocks and Bedford 2010; Horrocks and Rechtman 2009; Horrocks and Weisler 2006; Horrocks et al. 2004, 2012; Lentfer 2009; Loy 1994; Loy et al. 1992; Torrence, Wright and Conway 2004; Ussher 2009) with regard to starch and parenchyma within the Pacific. The analysis of a range of metric and descriptive nominal variables using light microscopy, Scanning Electron Microscopy (SEM), charting and multivariate statistics enabled a greater understanding of morphology, and therefore confidence in the identification of unknown archaeological samples. The multivariate statistical analysis identified those plant species or families that could be more easily discriminated from others, and highlighted where extra steps of visual checking would be required. There is still room for improvement, especially regarding the development of an automated system for starch classification with greater accuracy, measured through lowered rates of misclassification from multivariate statistical analysis. The introduction of new attributes and improved image analysis software may contribute to this (Coster and Field 2015).

It is hoped that these approaches to the extraction and identification of starch and vegetative storage parenchyma in this thesis will enable others engaged in the field of archaeobotany to collect and analyse these micro- and macrobotanical remains. This analysis is crucial towards elucidating the history of plant use in the Pacific region, where subsistence was and still is focussed primarily on the cultivation of starchy root, tuber and arboreal species. Starch and parenchyma derive directly from these organs which have been consumed in the past, and are still staple dietary components today. These provide clear evidence for diet and 268 subsistence, and therefore need to be further studied in terms of more detailed morphological and also biochemical analysis, as well as archaeobotanical applications. Each new piece of published research has built on the foundation of others, and this study is no exception. However, more work is yet to be done to understand starch preservation, contamination and biases that impact the interpretation of the archaeological record (Barton and Torrence 2015). This study has provided a baseline for the identification of assemblages from Tongatapu and these now need to be expanded to other locations in the Pacific through further research, transparency and consistent application of techniques.

Archaeobotanical research in Tonga and the Pacific The archaeobotanical analysis in this study was the first to have been conducted in the Tongan archipelago. However, the limited geographical and temporal scope of this analysis has meant that a chronology has been created from modelling high-resolution data from narrow time periods on one small area of an island within the archipelago. Time was a factor in the decision to restrict analysis to six small test units from three sites, but the primary focus of this project was to scale back analysis from large horizontal excavations that require coarse-grained analysis, instead utilising fine-grained archaeobotanical techniques that target extraction of large quantities of micro- and macrobotanical remains alongside faunal and artefactual material.

The successful recovery of these remains highlights the potential for further incorporation of these techniques within archaeological research strategies in tropical climates such as those on Tonga. A range of archaeological deposits were sampled, from dense midden to buried cultural deposits, and both starch and charred seeds, wood charcoal and endocarp were recovered from them all. The case study of charred parenchyma from Talasiu especially emphasises the nature of preservation of these fragile remains and points to the benefits of analysing them as a record of prehistoric plant use, or to complement microbotanical studies. The identification of charred remains was used in this study to both confirm and add to the lists of species cultivated and used within early production systems on Tongatapu. The parenchyma data added weight to the identification of some controversial or unexpected species at the late- Lapita site of Talasiu. This study has shown that it is crucial that this relatively unknown type of macro-remains be incorporated into future archaeobotanical studies that target diet and subsistence in the Pacific.

The three sites, from which archaeobotanical data was collected within this study, are well-dated and narrow in time depth, each representing only 100-200 years of occupation. This was seen as an advantage, enabling detailed data to be collected about subsistence within ‘snapshots’ of Tongan prehistory. Major inter-site variation was not expected, and was not observed, enabling initial modelling of singular archaeological production systems at each site. Inter-site variation, on the other hand, provided the basis for temporal comparison of these

269 systems and the timing of plant introductions into Tonga. These ‘snapshots’ of plant production and use from 2750-2650 cal BP, 1300-1000 cal BP, and 800-600 cal BP indicated the rate of change and interaction occurring during crucial periods in Tongan prehistory. Clearly, more research is required to fill the gaps, and elucidate in greater detail the role of plants in Tongan subsistence, and to test the hypothesis for agricultural development proposed here. Would the modelled systems, the basis for an alternative chronology, hold up under further testing using newly collected archaeobotanical data? There is only one way to find out and that is through further and more detailed archaeobotanical studies in the Tongan archipelago, especially at sites dated to early Lapita occupation and the later end of the Tongan sequence.

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Appendix A- Species in Reference Collection Marattiaceae Lecythidaceae Fabaceae Dioscoreaceae Dennstaedtiales Convolvulaceae Convolvulaceae Aspleniaceae Asparagaceae Arecaceae Araceae Apocynaceae Anacardiaceae Achariaceae Family Angiopteris Barringtonia racemosa Barringtonia asiatica Pueraria lobata fagifer Inocarpus Tacca leontopetaloide Dioscorea rotundata Dioscorea pentaphylla Dioscorea nummularia Dioscorea esculenta Dioscorea bulbifera Dioscorea alata Pteridium polpha Ipomoea batatas Ipomoea Asplenium Cordyline fruticosa nucifer Cocos sagittifolium Xanthosoma Epipremnum pinnatum merkusiiCyrtosperma Colocasia esculenta paeoniifolius Amorphophallus Alocasia macrorrhiza aurantiaca Tabernaemontana dulcis Spondias edule Pangium Species sp. sp. sp. a L. Reinw. Parkinson (Willd.) Ohwi (Willd.) (L.) (P.K. Latz.) (P.K. L. (Parkinson ex Zollinger) Fosberg Zollinger) ex (Parkinson (L.) Chev. (L.) (L.) (L.) Schott (L.) (L.) Poir. (L.) Kurz (L.) (L.) Schott (L.) s (L.) Kuntze s (L.) (L.) Schott (Hassk.) Lam. (L.) Engl. (L.) (L.) Spreng (L.) (L.) Schott (L.) (Denst.) Nicolson (Denst.) Gaudich. Giant ferns tree Powder-puff tree Fish-poison Kudzu chestnut Tahitian arrowroot Polynesian yam Guinea White yam Five-fingered yam Spiny yam Lesser yam Bitter Greater yam Bracken potato Giant sweet potato Sweet Spleenworts Ti Coconut Tannia Dragontail plant Giant swamp taro Taro lily Stink Giant taro none apple Otaheite fruit Football Common name halufe none futu aka ifi mahoa'a none lena palai ufilei hoi ufi aruhe none kumala none si niu talo futuna none via talo teve kape none vi none Local name (Tonga) or via via kana Madagascar to South Pacific Phillipines Oceans Indian/Pacific Asia Eastern/Southern Indo-Malaya/Melanesia Tropical Asia West Africa Asia Southeast Asia Southeast Asia Southeast World Old Tropics Asia Southeast Global Australia World New Oceans Indian/Pacific Himalayas to Australia Northern World Old Tropics Tropical America World Old Tropics Indo-Malaya Asia Southeast Madagascar to Indo-Malaya Guinea orTropical Asia New Tropical Asia/Pacific Indo-Malaya Asia Southeast Origins Alternative food Alternative soap Domestic- Foraging/hunting food Alternative food Alternative food Staple food Staple food Staple food Staple food Staple food Alternative Medicinal food Alternative food Alternative food Staple food Alternative thatch Domestic-fiber, food Alternative Medicinal timber thatch, vessels, fibre, Domestic- drink food, Staple food Staple food Alternative food Staple food Staple food Alternative food Staple Medicinal food Alternative food Alternative Utilisation

294

Zingiberaceae Solanaceae Rubiaceae Poaceae Piperaceae Pandanaceae Osmundiaceae Myristicaceae Musaceae Moraceae Family Zingiber zerumbet longa Curcuma tuberosum Solanum citrifoliaMorinda officinarum Saccharum Piper methysticum tectorius Pandanus Todea Horsfieldia palauensis Musa Ficus tinctorius Ficus copiosa heterophyllus Artocarpus altilisArtocarpus Species spp. (hybrids) spp. sp. Steud. G.Forst L. L. Fosberg (Parkinson) (L.) Roscoe ex Sm. Roscoe ex (L.) Forst. f. Parkinson L. Kaneh. L. Lam. Shampoo Shampoo ginger Turmeric Potato mulberry Indian Sugarcane Kava Screwpine fern King none Bananas/Plantains Dyer's fig Fig Jackfruit Breadfruit Common name angoango ango none nonu to kava fa/hingano none none fusi masi'ata none none mei Local name (Tonga) Indo-Malaya/Southern Asia Indo-Malaya/Southern Asia Southeast World New Asia Southeast Tropical Asia Vanuatu to Tropical Asia East Polynesia Zealand South to Africa New Palau Guinea Indo-Malaya/New India/Pacific Asia/Pacific Southeast Malaysia Guinea New Origins Medicinal Domestic-shampoo Domestic-dye food Staple Medicinal food Alternative food Staple Medicinal food/drug Alternative Medicinal thatch fibre, Domestic- food Alternative food Alternative food Alternative food Staple dye Domestic-fiber, food Alternative food Alternative food Staple food Staple Utilisation

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Appendix B- Description of Parenchyma Alocasia macrorrhiza (L.) G. Don: Corm (BG948)

Thin sections Epidermis- A layer of thick periderm protects a cortical region of around 20 radially organised rows of angular and elongated cells with very thin cell walls. Below this are two rows of similarly shaped cells with thicker walls that make up the vascular cambium.

Ground tissue- Interior to this is a wide region of amyliferous parenchymatous tissue through which run vascular bundles. The parenchyma cells tend to be between 80-130µm in length, and 70-100µm in width. Cells are mostly rounded and isodiametric in dimension (60%), however around 40% are elongated. Inter-cellular spaces are present but no cell contents were observed aside from cell nuclei. This was most likely a result of the process of the construction of the histological thin sections.

Vascular tissues- Vascular bundles are atactostele and so are widely separated and run apparently randomly through the organ. They are amphicribal concentric in arrangement with open ends (phloem surrounding the xylem apart from at each end of the bundle). The bundles range in length from 300-560µm, and 200-450µm in width.

Charred samples Charred dried- Within the cortex the vascular tissues preserve quite well. The cells within the conjunctive tissue become more irregularly rounded in shape but the dimensions are much the same as within the thin section. Cell walls become thicker and the inter-cellular spaces turn to solid charcoal. The tissues within the pith are altered by the formation of large cavities that compress the surrounding cells. Some starch granules preserve.

Charred fresh- The tissues are much the same as the sample charred from dried state, apart from the phloem which has become solid carbon within the cortex.

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Artocarpus altilis (Parkinson) Fosberg: Fruit (BG947)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- An outer epidermis composed of two to three layers of thickened radially organized rounded and isodiametric cells. Below this is a wide region of amyliferous parenchymatous tissue through which run vascular bundles. These parenchyma cells tend to be between 40-60µm in length, and 30-40µm in width. Cell shape and dimensions are relatively diverse, but the most common cell combination is rounded and isodiametric in dimension (40%), however a small number are angular and elongate (25%), or rounded and elongate (28%). Inter-cellular spaces are present and starch granules and druses were also observed within cells. Vascular bundles are atactostele and so are widely separated and run apparently randomly through the organ. The bundles are amphicribal u-shaped in arrangement (phloem surrounding the xylem in a u-shaped formation), and range in length from 100-600µm, and 70-280µm in width.

Charred samples Charred dried- Within the vascular tissues, the xylem preserved quite well but the phloem turned to solid carbon. The epidermis also becomes solid carbon along with many rows of cells within the ground tissue. The remaining cells within the conjunctive tissue are much the same as within the thin section.

Charred fresh- The tissues are generally preserved better within the freshly charred sample. The phloem has turned to solid carbon surrounding the xylem, but the ground and boundary tissues are still relatively the same morphologically.

Artocarpus altilis (Parkinson) Fosberg: Seed (BG947)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- An outer thin epidermis composed of only one layer of radially organised rounded and isodiametric cells with thickened cell walls. Interior to this is a wide region of amyliferous parenchymatous tissue through which run vascular bundles of varying sizes. The parenchyma cells tend to be between 60-80µm in length, and 50-65µm in 297 width. Cells are mostly rounded and isodiametric in dimension, however roughly 25% are elongated. Inter-cellular spaces are present and starch granules and druses were also observed within cells. The pith is composed of sclerenchyma. Vascular bundles are atactostele and so are widely separated and run apparently randomly through the organ. They are amphicribal concentric in arrangement (phloem surrounding the xylem). The bundles range in length from 80-900µm, and 75-360µm in width.

Charred samples Charred dried- Within the cortex the cells forming the conjunctive tissues have thicker cell walls and inter-cellular spaces become solid carbon. The cells within the pith are much the same as within the thin section. The xylem preserved well but the phloem turned to solid carbon during charring.

Charred fresh-Within the vascular tissues, the xylem preserved quite well but the phloem turned to solid carbon. The epidermis also becomes solid carbon and large cavities formed in the pith that compresses the surrounding cells. The remaining cells within the conjunctive tissue are much the same as within the fresh sample.

Angiopteris sp. :Rhizome (MP1152-005)

Thin sections Epidermis- An outer epidermis composed of around three layers of radially organised rounded and elongated cells with thickened cell walls.

Ground tissue- Interior to this is a region of cortical parenchymatous tissue. The parenchyma cells tend to be between 100-150µm in length, and 85-100µm in width. Cells are a combination of rounded and elongate in dimension (53%), and isodiametric (47%). Inter-cellular spaces are present and starch granules were also observed within cells. Within the parenchyma is a wide region of collenchyma with smaller rounded isodiametric cells and large inter-cellular spaces.

Vascular tissues- A dictyostele arrangement is present, however the vascular bundles could not be clearly observed within the histological thin sections.

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Charred samples Charred dried- The xylem preserves relatively well, but the surrounding phloem turns to solid carbon along with the fiber sheath. Most conjunctive parenchyma cells become solid carbon, but some small vessels survive within these. Some starch granules preserve.

Charred fresh-The vascular tissues are much the same as the dried charred sample. Many cells within the ground tissue flatten and compress, and large cavities forms within the pith. The epidermis becomes mostly solid carbon.

Asplenium sp.:Rhizome (EU003)

Thin sections Epidermis- An outer epidermis composed of one layers of radially organised angular and isodiametric cells with thickened cell walls. Below this is a broad region of approximately seven to eight rows of fibres that separate the epidermis and the conjuctive tissue within the pith.

Ground tissue- Interior to this is a region of cortical parenchymatous tissue. The parenchyma cells tend to be between 55-80µm in length, and 45-70µm in width. Cells are rounded in shape and predominantly isodiametric in dimension (77%), with a smaller percentage being elongate (23%). Inter-cellular spaces are present within this conjunctive tissue. Starch granules and druses were also observed within the cell contents.

Vascular tissues- A dictyostele overall arrangement is present, where the vascular chamber is broken into segments. The segments are amphicribal concentric in arrangement, where the xylem vessels are surrounded by the phloem. The sizes of these segments differ, with an total length range of 770-960µm and width range of 390-810µm.

Charred samples Charred dried and fresh- The xylem preserves relatively well, but the surrounding phloem turns to solid carbon along with the fiber sheath. Most conjunctive parenchyma cells become compressed and more angular, especially longitudinally, and some become solid carbon. Many very small starch grains survived the charring process.

299

Asplenium sp. :Rhizome (MP1152-009)

Thin sections Epidermis- An outer epidermis composed of one layers of radially organised angular and isodiametric cells with thickened cell walls. Below this is a broad region of fibres that separate the epidermis and the conjuctive tissue within the pith

Ground tissue- Interior to this is a region of amyliferous cortical parenchymatous tissue. The conjunctive parenchyma cells are relatively large compared to other ground tissue observed within the comparative collection. They range from 115-175µm in length, and 90-130µm in width. Cells are consistently angular in shape but range markedly in dimension. Around 53% are isodiametric and 47% are elongated. Inter-cellular spaces are present and starch granules were also observed within cells.

Vascular tissues- A dictyostele arrangement is present, where the vascular chamber is broken into segments. The segments are amphicribal concentric in arrangement, whereby the xylem is surrounded by the phloem, and range in length from 160-950µm and width from 110-570µm.

Charred samples Charred dried- The xylem preserves relatively well, but the surrounding phloem turns to solid carbon along with the fiber sheath. Tension fractures divide the conjunctive parenchyma cells which mostly turns to solid carbon, especially within the periderm.

Charred fresh- The vascular tissues are much the same as for the dried charred sample, however the ground tissue preserves much better in the fresh sample. The morphology of the cells is angular and broadly isodiametric. Some compression of cells is visible, and the periderm becomes mostly solid carbon. Fibre bundles also preserve.

300

Barringtonia asiatica (L.) Kurz: Fruit (EU2012-06)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- An outer epidermis composed of one layer of very small radially organised rounded and isodiametric cells with a thick outer cell wall. Below this is a row of palisade mesophyll cells. Interior to this is a wide region of amyliferous parenchymatous tissue. The parenchyma cells tend to be between 55-105µm in length, and 40- 75µm in width. Cells are consistently angular and isodiametric in dimension. Inter-cellular spaces are present but no cell contents were observable within the thin sections. Vascular bundles are of the eustele type, and amphivasal concentric in arrangement (xylem surrounding the phloem) with a fiber sheath. Bundles range in length from 85-160µm, and 85-150µm in width.

Barringtonia asiatica (L.) Kurz: Seed (EU2012-06)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- An outer epidermis composed of one layers of very small radially organised rounded and isodiametric cells with a thick outer cell wall. Below this is a region of cortical parenchyma, roughly the same size and shape as the ground tissue within the pith. Two rows of vascular cambium separates the cortex and pith. Within the cortical and conjunctive tissue within the pith, the parenchyma cells tend to be between 50-70µm in length, and 35-50µm in width. Cells are consistently mostly rounded and isodiametric in dimension, with roughly 33% being rounded and elongated. Inter-cellular spaces are present and starch was observable within the cells. Vascular bundles were not observed within the histological thin sections for this specimen.

Charred samples Charred dried- Cells within the epidermis, cortex and conjunctive tissue at the pith all survive charring with very little modification of morphology.

301

Charred fresh- These tissues do not preserve so well when charred from fresh state. The cells become mostly solid carbon within the cortex and pith, with a few small surviving cells interspersed throughout. Large cavities form within these regions. The epidermis becomes solid carbon with cell wall casts.

Barringtonia racemosa (L.) Spreng: Fruit (EU2012-12)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- A thin epidermis is composed of a single layer of angular and elongated cells with thick cell walls. Another possible ten rows of rounded and elongated cortical parenchyma below the epidermis. Within the conjunctive tissue within the pith, the parenchyma cells tend to be between 45-65µm in length, and 35-50µm in width. Cells are consistently mostly rounded and isodiametric in dimension, with roughly 28% being rounded and elongated. Inter-cellular spaces are present but no cell contents were observed. Vascular bundling is within a eustele arrangement typical of dicotyledons, and is of amphivasal concentric arrangement. The length range of the bundles is between 150-400µm, and the width is 110-300µm.

Barringtonia racemosa (L.) Spreng: Seed (EU2012-12)

302

Two rows of very small angular and elongated cells make up the vascular cambium which separates the cortex and pith. Within the conjunctive tissue within the pith, the parenchyma cells tend to be between 60-85µm in length, and 50-70µm in width. Cells are consistently angular and broadly isodiametric. Inter-cellular spaces are present and starch was observed within the parenchymatous cells. No vascular bundles were able to be observed within the histological thin sections.

Charred samples Charred dried- The parenchymous cells within the cortex and pith generally remain in good condition throughout the charring process. However the cells become more rounded, with thicker cell walls, and inter-cellular spaces become solid carbon. The epidermal cells become more compressed. Tension fractures form in the pith.

Colocasia esculenta (L.) Schott: Corm (BG1013)

Thin sections Epidermis-- An outer thickened epidermis composed of three layers. Below this is a region of around 20 rows of cortical parenchyma which is angular and elongated with thin cell walls. Two rows of angular and isodiametric cells with thick cell walls make up the vascular cambium which separates the cortex and pith.

Ground tissue- Within the conjunctive tissue within the pith, the amyliferous parenchyma cells are between 70-100µm in length, and 50-75µm in width. Cells are very inconsistent in shape and dimension. Around 33% of cells are rounded and isodiametric, 28% are rounded and elongated, 20% are angular and isodiametric, and another 20% are angular and elongated.. Inter- cellular spaces are present, and starch and raphides (calcium oxylate crystals) were observed within the cells contents.

303

Vascular tissues- Vascular bundles of both amphivasal concentric and u-shaped arrangement are present within an atactostele type of overall arrangement within the organ. The length range of the bundles is 230-580µm, and the width is 160-400µm.

Charred samples Charred dried- Within the vascular tissues, the xylem preserves very well but the phloem turns to solid carbon within the surrounding xylem. The cells walls within the conjunctive tissues of the pith become much thicker and the tissues become more vesicular. The epidermis is mostly turned to ash and shed. Where these cells do survive the preservation is very good and morphology does not differ much from the thin sections.

Charred fresh- The tissues charred from fresh state becomes more compressed, with inter- cellular spaces becoming solid carbon. Large cavities form within the pith. The vascular tissues survive in much the same form as for the dry charred sample.

Cordyline fruticosa (L.) A.Chev: Stem (EU006)

Thin sections Epidermis-- An outer thickened epidermis and below this is a region of cortical parenchyma which is rounded and isodiametric with thin cell walls. There is no obvious vascular cambium which separates the cortex and pith.

Ground tissue- Within the pith, the parenchyma cells are between 55-75µm in length, and 40- 55µm in width. Cells are consistently angular in shape but vary in dimension. Around 55% of cells are isodiametric and the remaining 45% are elongated. Inter-cellular spaces are present, but no cell contents were observed apart from the cell nuclei.

Vascular tissues- Vascular bundles of amphivasal concentric arrangement are present within an atactostele type of overall arrangement within the organ. The length range of the bundles is 200- 300µm, and the widths are between 100-200µm. Fibres also sheath each of the vascular bundles.

Charred samples Charred dried- The phloem within the vascular bundles becomes a cavity surrounded by xylem and fibres. Large cavities also form between these bundles in many places. Elsewhere the cells 304 become more compressed. The epidermis turns to solid carbon with casts of cell walls on the exterior surface.

Charred fresh- These tissues do not preserve so well when charred from fresh state. The cells become compressed, folded and flattened within the cortex and pith, with a few small surviving cells interspersed throughout. Pitting is visible on the inside of the cells. Large cavities form within these regions of conjunctive tissue. The epidermis becomes solid carbon with cell wall casts.

Cyrtosperma merkusii (Hassk.) Schott: Corm (EU2012- 09)

Thin sections Epidermis- Could not be viewed within the histological thin sections.

Ground tissue- The amyliferous parenchyma cells that form the conjunctive tissue are between 40-80µm in length, and 30-50µm in width. Cells are consistently rounded in shape but vary in dimension. Around 65% of cells are elongated and the remaining 35% are isodiametric. Inter- cellular spaces are present, and starch granules were observed within the cell contents.

Vascular tissues- Vascular bundles of amphivasal arrangement with open ends are present within an atactostele type of overall arrangement within the organ and so being present at random intervals and locations within the organ. The length range of the bundles is 300-500µm, and the widths are between 200-300µm.

Charred samples Charred dried- The phloem within the vascular bundles becomes compressed and solid carbon is some places, surrounded by xylem. The vascular bundles tend to break away from the surrounding tissue leaving a large cavity. Large cavities also form between the conjunctive tissues in many places. Elsewhere the cells are preserved well but the tissue becomes more vesicular overall.

Charred fresh- The fresh charred sample turned to ash.

305

Dioscorea alata (L.): Tuber (EU0013)

Thin sections Epidermis- An epidermis consisting of four layers of angular and elongated cells with thick cell walls.

Ground tissue- Below this is a region of amyliferous parenchyma cells that form the conjunctive tissue. These are between 80-110µm in length, and 50-75µm in width. Cells are consistently rounded in shape but vary in the regularity of shape and dimension. Approximately 28% of cells are irregularly rounded in shape and elongated in dimension. The remainder are rounded and elongated (45%) or isodiametric (29%). Inter-cellular spaces are present, and many starch granules were observed within the cell contents.

Vascular tissues- Vascular bundles are of closed collateral arrangement and are present within an atactostele type of overall arrangement within the organ and so being present at random intervals and locations within the organ. The length range of the bundles is 230-880µm, and the widths are between 85-340µm.

Charred samples Charred dried- The phloem within the vascular bundles becomes a cavity abutting the xylem within the collateral bundle. Large cavities also form within the pith. Elsewhere the cells become more irregularly shaped with thin walls. Many large starch granules are preserved within the cells.

Charred fresh- The cells within the cortex become mostly solid carbon that have amalgamated with inter-cellular spaces, with a few small cells interspersed. Large cavities form within these regions of conjunctive tissue. Starch is preserved with the cells. Vascular bundles are preserved in much the same form as for the sample that was dry charred.

306

Dioscorea bulbifera (L.): Aerial bulbil (BG942)

Thin sections Epidermis- One layer of thickened epidermis. Below this is a region of cortical parenchyma cells that are rounded and broadly isodiametric with thick cell walls. Many intercellular spaces are also present between cells. A region of vascular cambium separates the parenchyma within the cortex and the pith. This is made up of five rows of angular and elongated cells.

Ground tissue- The amyliferous parenchyma within the pith are between 95-130µm in length, and 75-100µm in width. Cells are consistently rounded in shape but vary in dimension. Approximately 62% of cells are rounded in shape and isodiametric in dimension. The remainder are rounded and elongated. Inter-cellular spaces are not present within the conjunctive tissue, but many starch granules were observed within the cell contents.

Vascular tissues- Vascular bundles could not be observed within the histological thin sections.

Charred samples Charred dried- The epidermis becomes solid carbon during charring. Cells within the conjunctive tissues of the pith remain rounded and isodiametric but the walls thicken and inter- cellular spaces become solid carbon. Large cavities form throughout the ground tissue.

Charred fresh- The boundary and ground tissues are mostly the same as for the sample charred from dried state, however tension fractures and larger cavities form.

Dioscorea esculenta (L.): Tuber (EU020)

Thin sections Epidermis- The epidermal region consists of three layers of angular and elongated cells.

Ground tissue- Below this is a region of amyliferous parenchyma which are between 150- 235µm in length, and 100-160µm in width. Cells are consistently rounded in shape but vary in dimension. Approximately 62% of cells are rounded in shape and isodiametric in dimension. The remainder are rounded and elongated. Inter-cellular spaces are sometimes present within the conjunctive tissue but are rare. Many compound starch granules were observed within the cell contents.

307

Vascular tissues- Vascular bundles are within an atactostele arrangement and so randomly run throughout the organ. The bundles themselves are of closed collateral arrangement, with no cambium between the vascular tissues. The length range of the bundles is 150-380µm, and the widths are between 80-240µm.

Charred samples Charred dried- The phloem within the vascular bundles becomes a cavity abutting the xylem within the collateral bundles. Large cavities also form between these bundles within the conjunctive tissue. Elsewhere the cells become shallower, and the epidermal cells become more compressed.

Charred fresh- These tissues do not preserve so well when charred from fresh state. The cells become more fragile with thinner cell walls. There is also carbonization of cell walls within large cavities that form in the ground tissue.

Dioscorea nummularia Lam.: Tuber (EU012)

Thin sections Epidermis- A thickened epidermis with a layer of five rows of angular and elongated cortical parenchyma cells.

Ground tissue- Below this is a region of amyliferous parenchyma which are between 75-115µm in length, and 60-100µm in width. Cells are consistently rounded in shape but vary slightly in dimension. Approximately 82% of cells are rounded in shape and isodiametric in dimension. The remainder are rounded and elongated. Inter-cellular spaces are present within the conjunctive tissue and starch granules were observed within the cell contents.

Vascular tissues- Vascular bundles are within an atactostele arrangement and so randomly run throughout the organ. The bundles themselves are of closed or open collateral arrangement, with a cambium separating the vascular tissues. The length range of the bundles is 155-435µm, and the widths are between 100-250µm.

Charred samples Charred dried- The parenchyma cells within the cortex become more irregularly shaped, while those in the pith mostly retain the original morphology. The vascular cambium becomes a 308 tension fracture and separates the cortex from the pith. Within the vascular tissues, the phloem either becomes a cavity or survives in original condition.

Charred fresh- The vascular and conjunctive tissues become more vesicular when charred from fresh state. The cell walls merge in places and become areas of solid carbon.

Epipremnum pinnatum (L.) Engl.: Corm (EU011)

Thin sections Epidermis- A thickened single-layer peridermis lies above a region consisting of around five rows of angular and elongated primary cortical parenchyma cells with thin cell walls. Another five rows of similarly shaped cells compose a region of secondary cortex. A row of vascular cambium that consists of one layer of rounded and elongated cells.

Ground tissue- Below this is a region of conjunctive tissue within the pith, consisting of amyliferous parenchyma cells which are between 70-95µm in length, and 50-70µm in width. Cells are consistently rounded in shape but vary slightly in dimension. Approximately 72% of cells are rounded in shape and isodiametric in dimension. The remainder are rounded and elongated. Inter-cellular spaces are present within this conjunctive tissue and small sparsely distributed starch granules were observed within the cell contents.

Vascular tissues- Vascular bundles are within a eustele arrangement. The bundles themselves are of amphivasal arrangement (phloem surrounded by xylem) in a u-shape pattern. The length range of the bundles is 220-375µm, and the widths are between 180-300µm.

Charred samples Charred dried and fresh- Cells within the boundary and conjunctive tissues become compressed and collapsed around the vascular bundles. Many crystals survive and can be seen within the cell contents. The xylem and fibre sheaths maintain their original morphology, but the phloem becomes a cavity within the surrounding xylem vessels.

309

Ficus copiosa Steud.: Fruit (EU-2012-13)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- Parenchyma cells are consistently rounded in shape and isodiametric in dimension. Inter-cellular spaces are visible between these. The length range of these cells is between 40-60µm, and the width range is between 30-45µm. Vascular bundles of closed collateral and amphicribal u-shaped or concentric form were observed within a eustele-type arrangement in the fruit. The bundles ranged in length from 70-380µm, and in width from 60-150µm.

Charred samples Charred dried and fresh- The parenchyma cells in the cortex of both the fresh and dry-charred samples become compressed and inter-cellular spaces become solid carbon. Seeds in the pith of the fruit survive charring and maintain original morphology. The surface morphology also retains the original ‘peaked’ texture. The ground tissues also become more vesicular overall. The vascular bundles retain the xylem but the phloem turns to solid carbon.

Ficus tinctoria G.Forst.: Fruit (MP1152-004)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- Parenchyma cells are consistently angular in shape but vary in dimension. The majority of cells are elongated in dimension (57%), with just under half being isodiametric (43%). Inter-cellular spaces are visible between these but no cell contents were recorded in this particular sample. The length range of these cells is between 55- 90µm, and the width range is between 25-70µm. No vascular bundles were recorded. 310

Charred samples Charred dried and fresh- The cells in the cortex become solid carbon within the wet-charred sample, but those in the dry-charred sample are still visible but have become compressed. The conjunctive tissues closer to the pith collapses during charring from both fresh and dried state. The seeds survive the charring process. Some original surface texture is also retained in parts.

Ipomoea batatas (L.) Lam.: Tuber (EU023)

Thin sections Epidermis- A region of periderm consisting of about four rows of radially-orientated angular and isodiametric cells with thin cell walls.

Vascular Tissues- Below this is a region of parenchymous secondary phloem outside the cambium consisting of angular cells that range from elongated (28%) to isodiametric (72%) in dimension. The length range of these cells is 80-115µm, and the widths vary from 60-90µm. The cambium consists of two rows of angular and elongated cells with slightly thicker walls, and separates the secondary phloem and xylem. The parenchyma within the secondary xylem themselves are the same as those within the phloem, however there are regions of anomalous tertiary growth adjacent to individual vessels. These parenchyma divide both periclinally and tangentially to produce concentric radiating rings of tertiary xylem.

Charred samples Charred dried- Parenchyma cells retain much of the original fresh morphology, but become possibly slightly shallower. Some tension fractures within the conjunctive tissues. The vascular tissues survive the charring process with no noticeable modification.

Charred fresh- The cells within the cortex become more compressed and often appear fractured and condensed. The epidermal cells retain original condition, but tension fractures separate these boundary tissues from the conjunctive tissues in places. Vascular tissues are much the same as within the dry-charred sample.

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Inocarpus fagifer (Parkinson ex Zollinger) Fosberg: Seed (BG955)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- A layer of endocarp protects the internal region of amyliferous parenchyma cells. This consists of a row of radially-orientated angular and isodiametric cells with very thick cell walls. The endocarp also divides the internal parenchymous tissue through the centre longitudinally. Contained within the bounds of the endocarp is a region of parenchymous tissue that consists of rounded and predominantly isodiametric cells (75%), with a length range of 45-100µm and width range of 35-75µm. The bundling arrangements of vascular tissues are amphicribal u-shaped and are within an atactostele arrangement whereby the tissues run seemingly randomly throughout the organ.

Charred samples Charred dried and fresh- The epidermal cells become solid carbon, and the cells within the conjunctive tissue become more condensed and compressed. Within the vascular bundles, the xylem preserves well but the phloem becomes solid carbon surrounding these tissues. Large vessels form in the pith of the organ.

Morinda citrifolia L.: Fruit (BG941)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- Parenchyma cells are consistently rounded in shape but vary in dimension. The majority of cells are isodiametric (85%), with a smaller percentage being elongated (15%). Inter-cellular spaces are visible between these but no cell 312 contents were recorded in this particular sample. The length range of these cells is between 40- 60µm, and the width range is between 30-50µm. Vascular tissues were visible but as this is not a sample of vegetative parenchyma these bundles did not match any of the typologies used for root or stem-derived tissues.

Charred samples Charred dried and fresh- Epidermal cells turn to solid carbon after charring from both dried and fresh states. The cells within the conjunctive tissues become compressed and cell walls can collapse. The xylem retains original condition, but the phloem becomes a cavity.

Musa sp.1: Fruit (BG1014)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- Within the conjunctive tissue, the cells are irregularly rounded in shape but vary in dimensions. The majority of cells are isodiametric (85%), with the remainder being more elongated (15%). Inter-cellular spaces are visible, as are large numbers of duct cavities located between the cells closer to the pith of the organ. The length range of these cells is 60-90µm, and the widths range from 50-70µm. Small vascular bundles are present in a eustele overall arrangement and are of amphicribal concentric arrangement. These bundles range in length from 80-115µm, and in width from 70-100µm. Many large starch granules are visible within the cell contents.

Charred samples Charred dried- Many large starch granules survive the charring process. Cells within the boundary tissues, cortex and pith all mostly collapse. The xylem and fibre bundles retain original morphology but phloem becomes a cavity.

Charred fresh- The parenchyma cells in the pith retain original condition in terms of shape and dimensions, but the inter-cellular spaces become cavities alongside the duct cavities. Tension fractures also form. The cortical region is more collapsed than the pith, with compressed and fractured cells walls.

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Musa sp.2: Fruit (BG995)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- The overall morphology of this sample is very similar to Musa sp.1 (BG1014) above, however there are several difference in cell size and shape that can allow differentiation between these two species. Cell lengths range from 73- 215µm, and widths range from 46-170µm. Within the conjunctive tissue, the cells are irregularly rounded in shape but vary in dimensions. The majority of cells are isodiametric (75%), with the remainder being more elongated (25%). Inter-cellular spaces are visible, as are large numbers of duct cavities located between the cells closer to the pith of the organ. Large vascular bundles are present in a eustele overall arrangement and are of amphicribal concentric arrangement. These bundles range in length from 220-680µm, and in width from 180-370µm. Bundles of fibres are disparately located within the organ. Many large starch granules are visible within the cell contents.

Charred samples Charred dried and fresh- Within both the samples wet and dry-charred the cell morphology retains original condition, however inter-cellular spaces become solid carbon. The duct cavities are also preserved. Some tension fractures form within the pith, and fractures also form along the vascular cambium. Overall the tissues become more vesicular. Within the vascular tissues, the phloem becomes a cavity.

Pandanus tectorius Parkinson: Fruit phalange (EU2011-03)

(Not vegetative storage parenchyma) 314

Thin sections Basic cell morphology and tissue arrangement- The epidermis is composed of one row of thickened, rounded and isodiametric cells. Below the epidermis is one row of similarly rounded and isodiametric cells with slightly thinner cell walls. This borders a region of parenchyma interspersed with vascular bundles surrounded by fibre sheaths. The vascular bundles are more highly concentrated towards the centre of the phalange and tend to be of amphicribal arrangement. The length range of these bundles is 260-750µm, and the width range is from 210- 430µm. The amyliferous parenchyma cells form the conjunctive tissue for the organ and are composed of rounded cells that are mostly isodiametric in dimension (63%). These cells range in length from 60-100µm and in width from 40-70µm. Inter-cellular spaces are often present between the cells.

Charred samples Charred dried and fresh- When charred, cell morphology becomes more irregularly rounded and slightly collapsed. The fibre sheaths surrounding the vascular bundles retain their original morphology along with the xylem, but the phloem becomes a cavity in the centre of the bundle.

Pangium edule Reinw.: Fruit (EU2012-02)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- A periderm forms the outer surface of the organ and this boundary tissue is made up of around six rows of angular and elongated cells. Below the periderm is a region of parenchyma interspersed with sclerenchyma. These cells have relatively thick cell walls. The conjunctive tissue is composed of rounded cells that are mostly isodiametric in dimension (75%). These cells range in length from 40-80µm and in width from 30-60µm. Inter-cellular spaces are often present between the cells. Xylem vessels are present near the pith of the fruit.

Charred samples Charred dried and fresh- The parenchmous cells within the ground and boundary tissues collapsed in many regions, especially closer to the pith. Other regions have retained the original

315 cell morphology. Within the vascular tissues the xylem vessels have also retained the condition of the fresh samples, but the phloem has become solid carbon.

Pueraria lobata (Willd): Root (BG974)

Thin sections Epidermis- A region of periderm consisting of about five rows of radially-orientated angular and isodiametric cells with thin cell walls.

Vascular tissues- Below this is a region of parenchymous cortex outside the cambium consisting of rounded cells that range from elongated (35%) to isodiametric (65%) in dimension. The length range of these cells is 25-45µm, and the widths vary from 20-30µm. The cambium separates the cortex from the stele, and is composed of an endodermis and pericyle of one row each. The parenchyma within the secondary xylem themselves are the same as those within the phloem, however there are regions of anomalous tertiary growth adjacent to individual vessels. There are many vessels within the xylem which is polyarch in arrangement. Many areas of tertiary xylem and phloem are also present and are dissected by medullary rays.

Charred samples Charred dried- Cells in all tissues become compressed, including the secondary xylem which also becomes fractured and compacted. However, the vessels within the xylem survive charring very well. Areas of solid carbon form around the stele and tension fractures separate the vascular tissues along the phloem in places.

Charred fresh- Within the sample charred from fresh state, the preservation of cell morphology within the stele was much greater. Large xylem vessels also maintain original condition and structure. The cambium split after a large tension fracture formed along this boundary, separating the secondary vascular tissues. Some starch granules also survived the charring process.

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Piper methysticum G.Forst: Root (EU-2011-02)

Thin sections Epidermis- A region of periderm consisting of three rows of angular and isodiametric cells that are thick walled and radially orientated.

Vascular tissues- Below this is cortical amyliferous parenchymous tissue within the primary tissues, made up of angular and isodiametric cells with very few inter-cellular spaces. Within this region are wide medullary ray sections made up of ligneous fibres which are thick walled cells rounded in shape in transverse section, and also xylem vessels. Scanty paratrachial to vasicentric layers of xylem cells abut these rays and are differentiated by thinner walls and more angular shape. Bundles of phloem are contained within these areas of xylem. Exterior to the cambium are further bundles of xylem within the parenchymous tissues. At the centre of the root is a region of pith. The cells within the pith have a length range of 45-75 µm and a width range of 35-60µm, and are rounded and isodiametric in dimension with many inter-cellular spaces.

Charred samples Charred dried and fresh- When charred from both fresh and dried state, the root of the kava plant retains much of the original ground and vascular tissue morphology. Cell shape and dimensions do not change drastically, however some small cavities can form within the medullary rays of fibres alongside the xylem vessels. Large amounts of starch survive the charring process within the ground tissues between the rays of fibres, in fact these cells are jam- packed full of starch.

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Pteridium sp.: Rhizome (EU002)

Thin sections Epidermis- An outer epidermis composed of around three layers of radially organised rounded and elongated cells with thickened cell walls.

Ground tissue- Interior to this is a region of cortical parenchymatous tissue. The parenchyma cells tend to be between 80-115µm in length, and 65-90µm in width. Cells are a combination of angular in shape and elongate in dimension (60%), and isodiametric (40%). Inter-cellular spaces are not present and starch granules were also observed within cells.

Vascular tissues- A dictyostele arrangement is present, however the vascular bundles could not be clearly observed within the histological thin sections.

Charred samples Charred dried and fresh- Most tissues become either very compacted or turn to solid carbon when charred from either fresh or dried state. Large cavities and tension fractures form throughout the ground and vascular tissues. This species within the genus Pteridium would be very hard to identify as charcoal.

Solanum tuberosum (L.): Stem tuber

Thin sections Epidermis- An outer thickened epidermis outside a region of cortical parenchyma. The cortex consists of about four to five rows of angular and elongate cells with thinner walls. The vascular cambium is also two layers of angular cells that are slightly more isodiametric

Ground tissue- Interior to this is a region of amyliferous parenchymatous tissue. The parenchyma cells tend to be between 90-130µm in length, and 70-100µm in width. Cells are irregularly rounded in shape and elongated in dimension (53%), and isodiametric (47%). Inter- cellular spaces are not present but many large starch granules were observed within cells.

Vascular tissues- Vascular bundles are bicollateral in arrangement and tend to be long and thin in overall dimensions. The bundles themselves are arranged within an atactostele orientation, and so appear to be randomly located within the organ.

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Spondias dulcis (L.): Fruit (BG981)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- A region of periderm forms the boundary tissue for this fruit, composed of an outer thickened periderm, and a cortical region of approximately five to ten rows of rounded and isodiametric cells with thick cell walls. The thickness of the periderm varies around the circumference of the organ. The ground tissue within the pith has thinner cell walls and is angular in shape and primarily elongated in dimension (75%). These cells within the conjunctive tissue range in length from 85-160µm , and in width from 55-95µm. There are no inter-cellular spaces visible within the cells, but many starch granules can be seen within the cell contents. Vascular bundles vary between closed collateral and amphicribal concentric arrangement within an overall eustele arrangement of these tissues within the organ. These bundles range in length from 280-400µm, and in width from 140-240µm.

Charred samples Charred dried- The tissues within the periderm and cortex becomes solid carbon with small vessels interspersed throughout these regions. Other ground tissues generally maintain the original cell shape but these can also appear fractured. Within the vascular tissues, the xylem preserves well but is surrounded by a band of phloem that has turned to solid carbon. Large cavities form in the pith.

Charred fresh-The tissues within the fresh-charred sample have many of the same modifications as within the sample dried prior to charring. The periderm and cortex becomes a region of solid carbon, and some ground tissues within the pith retain the original morphology with angular and elongated cells. However aside from these areas of cells, the pith mostly becomes highly vesicular with large cavities and tension fractures.

319

Syzygium malaccense (L.)Merr.: Fruit (EU2012-01)

(Not vegetative storage parenchyma) Thin sections Basic cell morphology and tissue arrangement- A thin epidermis composed of a single row of angular and elongated cells, with a thicker exterior wall. Several rows of small cells that are angular in shape and more isodiametric in dimension are below this within the cortex, followed by a region of parenchmous ground tissues. The ground tissue is composed of larger angular and mostly elongated cells (55%), and very few inter-cellular spaces. The length range of these cells is from 75-120µm, and the width range is from 45-85µm. Vascular bundles are dispersed throughout the cortex and pith within a eustele arrangement, and are of amphicribal ‘stellate’ arrangement themselves. Within these bundles the xylem forms a stellate polyarch pattern with the many steles, surrounded by the phloem. These bundles range is length from 250-620µm, and width from 150-330µm.

Charred samples Charred dried and fresh- All ground tissues collapse and become fractured, or meld into solid carbon during the charring process. Within the vascular tissues, the phloem surrounding the xylem with the stele turns to solid carbon but the xylem maintains its original morphology. Large cavities form within the ground tissue and tension fractures separate the areas of solid carbon from each other, creating a highly vesicular overall appearance.

Tabernaemontana aurantiaca Gaudich.: Fruit (EU2012-05)

(Not vegetative storage parenchyma)

320

Thin sections Basic cell morphology and tissue arrangement- The epidermis a single row of tangentially flattened angular and elongated cells. Interior to this is a region of amyliferous parenchymatous tissue. These conjunctive parenchyma cells tend to be between 35-60µm in length, and 25- 40µm in width. Cells are rounded in shape and elongated in dimension (53%), and isodiametric (47%). Inter-cellular spaces are present but no cell contents were visible within the cell walls. Vascular bundles are within an eustele arrangement and tend to be amphicribal concentric in arrangement. The length range of the bundles is 175-370µm, and the range of widths is between 100-230µm.

Charred samples Charred dried and fresh- The epidermal cells become solid carbon during the charring process from both dry and fresh states. Fibre bundles survive in original condition. The ground tissues become collapsed, solid carbon in places and generally more vesicular in appearance. Vascular tissues were not visible as such within charcoal.

Tacca leontopetaloides (L.) Kuntz: Root tuber (EU2015)

Thin sections Epidermis- The periderm consists of three rows of radially orientated angular and elongated cells with thin cell walls, with the outermost row having a thicker external wall.

Ground tissue- Below this is a region of conjunctive tissue within the pith, consisting of amyliferous parenchyma cells which are between 70-100µm in length, and 60-80µm in width. Cells are consistently rounded in shape but vary slightly in dimension. Approximately 83% of cells are rounded in shape and isodiametric in dimension. The remainder are rounded and elongated. Many inter-cellular spaces are present within this conjunctive tissue and numerous starch granules, druses and calcium oxylate crystals are visible within the cell contents.

Vascular tissues- Vascular bundles are within an atactostele arrangement and tend to be bicollateral in arrangement. The length range of the bundles is 250-530µm, and the range of widths is between 100-270µm.

321

Charred samples Charred dried- In general cell morphology is preserved in original condition within the ground and boundary tissues. Some small cavities form within the pith and cortex, along with a small number of tension fractures nearer the pith. Bundles of raphides survive the charring process. Within the vascular tissues the phloem becomes a cavity, but the xylem preserves well.

Charred fresh- Many large cavities form within the pith of the sample charred from fresh state. The interior of these cavities have the outline of cell walls on the surface. Ground tissues become more vesicular overall, and the epidermal cells become compressed in shape. Again small bundles of raphides survive within cells in the ground tissues.

Todea sp.: Rhizome (EU004)

Thin sections Epidermis- An outer epidermis composed of one layers of radially organised angular and isodiametric cells with thickened cell walls. Below this is a broad region of fibres that separate the epidermis and the conjuctive tissue within the pith

Ground tissue- Interior to this is a region of cortical parenchymatous tissue. The parenchyma cells tend to be between 75-135µm in length, and 60-105µm in width. Cells are angular in shape and predominantly isodiametric in dimension (65%), with a smaller percentage being elongate (35%). Inter-cellular spaces are not present and starch granules were also observed within cells.

Vascular tissues- A dictyostele arrangement is present, where the vascular chamber is broken into segments.

Charred samples Charred dried- The ground tissue preserves the charring process in original condition, but the boundary tissue becomes solid carbon. Within the dictyostele vascular bundles, the xylem survives however the surrounding phloem becomes solid carbon.

Charred fresh- The charred ground tissue within the fresh-charred sample are similarly in good original condition. The epidermis and regions of sclerenchyma also mostly retain their morphology, with some areas of more compressed cells. Bundles of fibres also preserve. The

322 phloem becomes a large cavity within the vascular bundles, leaving only some areas of xylem cells within the vascular chamber.

Xanthosoma sagittifolium (L.) Schott.: Corm (EU-2012- 10)

Thin sections Epidermis- The periderm consists of four rows of radially orientated angular and isodiametric cells with thin cell walls, with the outermost row having a thicker external wall.

Ground tissue-. The amyliferous parenchyma cells that form the conjunctive tissue are between 55-80µm in length, and 50-60µm in width. Cells are consistently rounded in shape and isodiametric in dimension. Inter-cellular spaces are present, and many compound starch granules were observed within the cell contents.

Vascular tissues- Vascular bundles are within an atactostele arrangement and tend to be amphivasal concentric in arrangement (phloem surrounded by xylem). The length range of the bundles is 90-270µm, and the range of widths is between 80-230µm.

Charred samples Charred dried- Ground tissues survive mostly in original condition, however these become more vesicular. Bundles of raphides are present within cavities between cells. Tension fractures form along the vascular cambium, separating the boundary tissues from the ground tissues. The peridem becomes compressed.

Charred fresh- There is much greater modification of tissues within the sample charred from fresh state. Areas of ground tissue have survived in original condition, but the majority of these tissues become solid carbon with many small vesicles along with the periderm. Large tension fractures and cavities form withi the pith with the outlines of cell walls on the interior surfaces. Within the vascular bundles, the xylem survives but the phloem becomes solid carbon.

323

Zingiber sp.: Rhizome (BG957)

Thin sections Epidermis- A thickened periderm composes the surface of the rhizome, with around ten rows of thick walled cells that are rounded and broadly isodiametric in dimension.

Ground tissue-. The amyliferous parenchyma cells that form the conjunctive tissue below the periderm are between 55-80µm in length, and 50-60µm in width. Cells are consistently irregularly rounded in shape but vary in dimension. The majority of cells are elongated in dimension (60%) but a small number are broadly isodiametric (40%). Inter-cellular spaces are present, but no cell contents were visible during light microscopy of the histological thin sections.

Vascular tissues- Vascular bundles are within an eustele arrangement and tend to be closed collateral in arrangement, with a fibre sheath surrounding the bundles. The bundles are more concentrated in the pith of the rhizome, and more disparate towards the periderm. These range in length from 296-447µm and width from 240-352µm.

Charred samples Charred dried and fresh- The vascular bundles are easily identifiable as such within the charred samples, with the fibre sheath and xylem tissues surviving the charring process in good original condition. The phloem does not survive and becomes a cavity within these bundles. The surrounding conjunctive tissue collapses and becomes more irregularly shaped with thin cells walls. Cells within the periderm also become more compressed.

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Zingiber sp.: Rhizome (EU007)

Thin sections Epidermis- The periderm consists of around ten rows of rounded and elongated thick walled cells that are radially orientated. The exterior wall of the outermost layer is thicker than the other cell walls. There is a vascular cambium below this that is a single layer of more angular and elongated cells.

Ground tissue-. The pith of the organ is composed of conjunctive amyliferous parenchyma cells that are consistently rounded in shape, and predominantly isodiametric in dimension (60%). This differs from another unidentified species from the Zingiberaceae family that has been included within the comparative collection and is described above, where a smaller percentage of cells are isodiametric. The cells range in length from 75-105µm, and 50-85µm in width. Inter-cellular spaces are present.

Vascular tissues- Vascular bundles are within a eustele arrangement and tend to be amphicribal concentric in arrangement, with a fibre sheath surrounding the bundles. The bundles are more concentrated in the pith of the rhizome, and more disparate towards the periderm. These range in length from 160-309µm and width from 112-240µm.

Charred samples Charred dried and fresh- The xylem within the vascular bundles survive the charring process in good original condition, however the phloem and fibre sheath does not survive and become solid carbon surrounding these bundles. The surrounding conjunctive tissue mostly becomes compressed but there are regions of surviving cells with original morphology between these bands of compressed cells. Cells within the periderm become solid carbon with small vesicles.

325

Appendix C- Starch Images

Alocasia macrorrhiza (L.) G.Don: Corm (BG948)

Amorphophallus paeoniifolius (Dennst) Nicolson: Corm (YEN)

Artocarpus altilis (Parkinson) Fosberg: Fruit (BG947)

Barringtonia asiatica (L.) Kurz: Fruit (EU2012-06)

326

Colocasia esculenta (L.) Schott: Corm (BG1013)

Curcuma longa L.: Rhizome (EU-2008-01)

Cyrtosperma merkusii (Hassk.) Schott: Corm (EU2012-09)

Dioscorea alata (L.): Tuber (EU0013)

Dioscorea bulbifera (L.): Aerial bulbil (BG942)

327

Dioscorea esculenta (L.): Tuber (EU020)

Dioscorea nummularia Lam.: Tuber (EU012)

Dioscorea pentaphylla (L.): Tuber (YEN)

Inocarpus fagifer (Parkinson ex Zollinger) Fosberg: Seed (BG955)

Ipomoea batatas (L.) Lam.: Tuber (EU023)

328

Ipomoea polpha R.W Johnson: Tuber (YEN)

Morinda citrifolia L.: Fruit (BG941)

Musa sp.1: Fruit (BG1014)

Musa sp.2: Fruit (BG995)

Piper methysticum G.Forst: Root (EU-2011-02)

329

Solanum tuberosum (L.): Stem tuber

Spondias dulcis (L.): Fruit (BG981)

Tacca leontopetaloides (L.) Kuntz: Root tuber (EU2015)

Xanthosoma sagittifolium (L.) Schott.: Corm (EU-2012-10)

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Appendix D- Archaeobotanical Research in the Pacific Mid Holocene- Mid LapitaHolocene- Pleistocene/Early Holocene Period Cycadaceae Combretaceae Calophyllaceae Burseraceae Boraginaceae Arecaceae Araceae Anacardiaceae Achariaceae Sapindaceae Pandanaceae Musaceae Dioscoreaceae Euphorbiaceae Burseraceae Araceae Achariaceae Family Cycas circinalisCycas Terminalia catappa inophyllum Calophyllum spp. Canarium Cordia sp. nucifera Cocos merkusii Cyrtosperma Colocasia esculenta dulcis Spondias Dracontomelon sp. edule Pangium Pometia pinnata sp. Pandanus (Eumusa/Australimusa) sp. Musa Dioscorea sp. Dioscorea cf.hispida Dioscorea alata Aleurites moluccana spp. Canarium Colocasia esculenta Alocasia macrorrhiza merkusii Alocasia longiloba/Cyrtosperma edule Pangium Taxa Arawe Is, PNG Is, Arawe Mussau PNG Is, PNG Is, Arawe Mussau PNG Is, Mussau PNG Is, PNG Is, Arawe Mussau PNG Is, PNG Is, Arawe Mussau PNG Is, PNG Is, Arawe Ngerchau, Palau Urupiv, Vanuatu Samoa Mulifanua, Solomons Islands, Reef PNG Islands, Anir Fiji Bourewa, Mussau PNG Is, Mussau PNG Is, PNG Is, Arawe Mussau PNG Is, PNG Dongan Sepik-Ramu, Cave, PNG Dongan Sepik-Ramu, Cave, PNG Swamp, Kuk PNG Swamp, Kuk Malaysia Cave, Niah Malaysia Cave, Niah PNG Dongan Sepik-Ramu, Cave, PNG Dongan Sepik-Ramu, Cave, Lake Wanum, PNG PNG Swamp, Kuk Malaysia Cave, Niah Is. Solomon Island, Cave,Buka Kilu Is. Solomon Island, Cave,Buka Kilu Malaysia Cave, Niah PNG Dongan Sepik-Ramu, Cave, Location Seed Seed Seed Seed Endocarp Endocarp Seed Seed Endocarp Endocarp Pollen Starch,xylem raphides, Starch Starch Starch Starch,xylem raphides, Endocarp Seed Seed Pericarp Seed Drupes Phytoliths Starch Parenchyma Starch Seed Seed Pollen Starch Parenchyma Starch Starch Starch Seed Botanical remains Matthews and Gosden 1997 Kirch 1987,1988, 1989 Matthews and Gosden 1997 Kirch 1987,1988, 1989 Kirch 1987,1988, 1989 Matthews and Gosden 1997 Kirch 1987,1988, 1989 Matthews and Gosden 1997 Kirch 1987,1988, 1989 Matthews and Gosden 1997 and WardAthens 2001 Horrocks 2013al. et Horrocks and 2004;Bedford Crowther 2009 Crowther 2009 Crowther 2001, 205, 2009 Horrocks and 2007Nunn Kirch 1987,1988, 1989 Kirch 1987,1988, 1989 Matthews and Gosden 1997 Kirch 1987,1988, 1989 1991al. et Swadling 1991al. et Swadling 1985Wilson 2006;al. et Fullagar 2003,Denham al. et 2004; Denham 2007 Paz and Barton 2007 Paz and Barton 2007 1991al. et Swadling 1991al. et Swadling 1995Haberle 2003Denham al. et Paz and Barton 2007 Loy 1992al. et Loy 1992al. et Paz and Barton 2007 1991al. et Swadling Publication 3840-1580 BP 3200-200 0 BP 3840-1580 BP 3200-200 0 BP 3200-200 0 BP 3840-1580 BP 3200-200 0 BP 3840-1580 BP 3200-2000 BP 3840-1580 BP 4500 BP 2700 BP c. 2750 BP c. 3100 BP c.3300 BP 3050-2500 BP 3200-200 0 BP 3200-200 0 BP 3840-1580 BP 3200-200 0 BP >5,500 BP >5,500 BP 10,220-6440 cal. BP 10,220- 9910 cal. BP 21,130 cal. BP <40,000 BP >5,500 BP >5,500 BP ~ 9,000 BP 10,220-6440 cal. BP 23,850-23,020 cal. BP 28,000 cal. BP 28,000 cal. BP <40,000 BP >5,500 BP Dates

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Period Family Taxa Location Botanical remains Publication Dates Mussau Is, PNG Seed Kirch 1987,1988, 1989 3200-200 0 BP Dioscoreaceae Dioscorea esculenta Bourewa, Fiji Starch, raphides, xylem Horrocks and Nunn 2007 3050-2500 BP Vao and Urupiv, Vanuatu Starch Horrocks et al. 2013 3000-2600 BP Dioscorea nummularia Vao, Vanuatu Starch Horrocks and Bedford 2010 3100-2800 BP Dioscorea pentaphylla Vao, Vanuatu Starch Horrocks and Bedford 2010 3100-2800 BP Euphorbiaceae Aleurites moluccana Arawe Is, PNG Seed Matthews and Gosden 1997 3840-1580 BP Fabaceae Inocarpus fagifer Mussau Is, PNG Pericarp Kirch 1987,1988, 1989 3200-2000 BP Musaceae Musa sp. Reef Islands, Solomons Starch, phytoliths Crowther 2009 c.3100 BP Urupiv, Vanuatu Phytoliths Horrocks and Bedford 2011 3000-2700 BP Matilau, Vanuatu Phytoliths Horrocks et al. 2009 2800-2500 BP Pandanaceae Pandanus spp. Arawe Is, PNG Drupes Matthews and Gosden 1997 3840-1580 BP Mussau Is, PNG Drupes Kirch 1987,1988, 1989 3200-200 0 BP Sapindaceae Pometia pinnata Mussau Is, PNG Seed Kirch 1987,1988, 1989 3200-200 0 BP Late Holocene Araceae Alocasia macrorrhiza Katem compound, Kosrae Parenchyma Athens et al. 1996 c.1900 BP Futuna Pollen Piazza and Frimigacci 1991 ~600 BP Colocasia esculenta Me Aure Cave, New Caledonia Starch Horrocks et al. 2008 2700-1800 BP Avai'o'vuna Swamp, Vavau, Tonga Pollen Fall 2005; Fall 2010 ca. 2600 BP Ngofe Marsh, Vavau, Tonga Pollen Fall 2010 ca. 2000 BP Finpea, Kosrae Pollen Athens et al. 1996 1523-1350 BP Tangatatau, Mangaia Parenchyma Kirch et al. 1995 788-331 BP Mauna Kea Adze Quarry, Hawaii Parenchyma Allen 1981, 1984 ~775 BP La Perouse Bay, Rapanui Starch Cummings 1998 650-150 BP Te Niu, Rapanui Starch Horrocks and Wozniak 2008 650-150 BP Stonefields, Auckland Starch, xylem Horrocks and Lawlor 2006 550-380 cal. BP Nuku Hiva, Marquesas Starch Allen and Ussher 2013 550-350 BP Rangihoua Bay, NZ Starch Horrocks et al. 2004 ~500 BP Triangle Flat, NZ Starch Horrocks et al. 2004 430-260 BP Rano Kau, Rapanui Starch Horrocks et al. 2012 post. 1605-1414 cal. BP Aspouri Peninsula, NZ Starch, xylem Horrocks et al. 2007 Undated Motutangi, NZ Starch Horrocks and Barber 2005 Undated Anaura Bay, NZ Starch Horrocks et al. 2008b Undated Pitcairn Island Starch, xylem Horrocks and Weisler 2006 Undated Cyrtosperma merkusii Tafunsak, Kosrae Pollen Athens et al. 1996 1997-1709 BP Finpea, Kosrae Pollen Athens et al. 1996 1523-1350 BP Henderson Island, Pitcairn Group Leaf parenchyma Hather and Weisler 2000 post. 950 BP Tangatatau, Mangaia Parenchyma Kirch et al. 1995 788-331 BP Arecaceae Cocos nucifera Katem compound, Kosrae Endocarp Athens et al. 1996 c.1900 BP Tangatatau, Mangaia Endocarp/mesocarp Kirch et al. 1995 788-331 BP Mauna Kea Adze Quarry, Hawaii Endocarp Allen 1981, 1984 ~775 BP Nikunau, Kiribati Endocarp Piazza 1998 520-305 BP Tiwaka Valley, New Caledonia Wood Dotte-Sarout et al. 2013 post. 650 BP Kaho'olawe/Kuli'ou'ou/Lapakahi/ Anaeho'omalu/Kane'ohe/Kalahuipua'a/ Ka'ahumanu, Hawaii Endocarp, husk Allen 1984 Varied 332

Period Family Taxa Location Botanical remains Publication Dates Asparagaceae Cordyline fruticosa Avai'o'vuna Swamp, Vavau, Tonga Pollen Fall 2005; Fall 2010 ca. 2600 BP Ngofe Marsh, Vavau, Tonga Pollen Fall 2010 ca. 2200 BP Katem compound, Kosrae Wood Athens et al. 1996 c.1900 BP Tangarutu, Rapa Wood Prebble and Anderson 2012 900-320 BP Tangatatau, Mangaia Wood Kirch et al. 1995 788-331 BP Kahikinui, Hawaii Wood Kirch et al. 2005 460-285 cal. BP Kaho'olawe/Kuli'ou'ou/Kalahuipua'a, Hawaii Wood Allen 1984 Varied Boraginaceae Cordia subcordata. Mauna Kea Adze Quarry, Hawaii Wood Allen 1981, 1984 ~775 BP Nikunau, Kiribati Wood Piazza 1998 520-305 BP Tiwaka Valley, New Caledonia Wood Dotte-Sarout et al. 2013 post. 650 BP Nuku Hiva, Marquesas Wood Huebert et al. 2010 500-300 BP Burseraceae Canarium spp. Kalahuipua'a, Hawaii Endocarp Allen 1984 700-200 BP Calophyllaceae Calophyllum inophyllum Tiwaka Valley, New Caledonia Wood Dotte-Sarout et al. 2013 post. 650 BP Combretaceae Terminalia catappa Katem compound, Kosrae Parenchyma Athens et al. 1996 c.1900 BP Tiwaka Valley, New Caledonia Wood Dotte-Sarout et al. 2013 post. 650 BP Convovulaceae Ipomoea batatas Tangatatau, Mangaia Parenchyma Hather and Kirch 1991 ~950 BP Nuku Hiva, Marquesas Starch Allen and Ussher 2013 750-350 BP Whangapoua, NZ Starch Horrocks et al. 2007 704-550 cal. BP Kona, Hawaii Starch, xylem Horrocks and Rechtman 2009 650-325 BP La Perouse Bay, Rapanui Pollen Cummings 1998 650-150 BP Te Niu, Rapanui Starch Horrocks and Wozniak 2008 650-150 BP Stonefields, Auckland Starch Horrocks and Lawlor 2006 550-380 cal. BP Lapakahi, Hawaii Parenchyma Allen 1984; Rosendahl 1972 525-225 BP Harataonga Bay, NZ Starch Horrocks et al. 2002 562-386 cal. BP Rangihoua Bay, NZ Starch Horrocks et al. 2004 ~500 BP Triangle Flat, NZ Starch Horrocks et al. 2004 430-260 BP Rano Kau, Rapanui Starch Horrocks et al. 2012 post. 1605-1414 cal. BP Hamurana Rd, Bay of Plenty, NZ Starch Horrocks et al. 2003 260-160 BP Pouerua/Puketona, NZ Starch, xylem Horrocks et al. 2004 pre. 150 BP Avai'o'vuna Swamp, Vavau, Tonga Pollen Fall 2005; Fall 2010 Historic Motutangi, NZ Starch Horrocks and Barber 2005 Undated Anaura Bay, NZ Starch Horrocks et al. 2008b Undated Pitcairn Island Starch, xylem Horrocks and Weisler 2006 Undated Cucurbitaceae Lagenaria siceraria Rano Kau, Rapanui Pollen Horrocks et al. 2012 post. 1605-1414 cal. BP Tangarutu, Rapa Pericarp Prebble and Anderson 2012 900-320 BP Te Niu, Rapanui Pollen Horrocks and Wozniak 2008 650-150 BP Harataonga Bay, NZ Pollen Horrocks et al. 2002 562-386 cal. BP Pouerua/Puketona, NZ Pollen Horrocks et al. 2004 pre. 150 BP Kaho'olawe/Kuli'ou'ou/ Anaeho'omalu/Kane'ohe/Kalahuipua'a/ Ka'ahumanu, Hawaii Endocarp Allen 1984 Varied 333

Period Family Taxa Location Botanical remains Publication Dates Dioscoreaceae Dioscorea alata Halawa, Hawaii Parenchyma Allen 1984 750-450 BP Te Niu, Rapanui Starch Horrocks and Wozniak 2008 650-150 BP Anaura Bay, NZ Starch Horrocks et al. 2008b Undated Dioscorea esculenta Me Aure Cave, New Caledonia Starch Horrocks et al. 2008 2700-1800 BP Dioscorea sp. Me Aure Cave, New Caledonia Starch Horrocks et al. 2008 2700-1800 BP Rano Kau, Rapanui Starch Horrocks et al. 2012 post. 1605-1414 cal. BP Nuku Hiva, Marquesas Starch Allen and Ussher 2013 550-350 BP Motutangi, NZ Starch Horrocks and Barber 2005 Undated Hather 1994b; Green and Upolu, Samoa Parenchyma Davidson 1969, 1974 900-540 BP Euphorbiaceae Aleurites moluccana Kaho'olawe/Kuli'ou'ou/Lapakahi/ Kane'ohe/Halawa/Kalahuipua'a/Anahulu/Ka'ahumanu, Hawaii Endocarp, seed Allen 1984 Varied Tangarutu, Rapa Endocarp Prebble and Anderson 2012 900-320 BP Tangatatau, Mangaia Endocarp Kirch et al. 1995 788-331 BP Fabaceae Inocarpus fagifer Katem compound, Kosrae Parenchyma Athens et al. 1996 c.1900 BP Nuku Hiva, Marquesas Wood Huebert 2014 post. 1700 Moraceae Artocarpus altilis Katem compound, Kosrae Parenchyma Athens et al. 1996 c.1900 BP Finpea, Kosrae Pollen Athens et al. 1996 1523-1350 BP Malsu, Kosrae Pollen Athens et al. 1996 1264-1150 BP Papeno'o Valley, Society Islands Wood Orliac 1997 ~650 Tiwaka Valley, New Caledonia Wood Dotte-Sarout et al. 2013 post. 650 BP Nuku Hiva, Marquesas Starch Allen and Ussher 2013 550-350 BP Nuku Hiva, Marquesas Wood Huebert 2014 682 BP-present Opunohu Valley, Society Islands Exocarp Kahn and Ragone 2013 ~250 BP cf. Artocarpus altilis Futuna Pollen Piazza and Frimigacci 1991 ~600 BP Broussonetia papyrifera Rangihoua Bay, NZ Pollen, phytoliths Horrocks et al. 2004 ~500 BP Rano Kau, Rapanui Pollen Horrocks et al. 2012 post. 1605-1414 cal. BP Ficus sp. Tiwaka Valley, New Caledonia Wood Dotte-Sarout et al. 2013 post. 650 BP Musaceae Musa sp. Rano Kau, Rapanui Phytoliths Horrocks et al. 2012 post. 1605-1414 cal. BP Tangatatau, Mangaia Leaf parenchyma Kirch et al. 1995 788-331 BP Kona, Hawaii Phytoliths Horrocks and Rechtman 2009 650-325 BP Futuna Pollen Piazza and Frimigacci 1991 ~600 BP Kaho'olawe, Hawaii Wood Allen 1984; McAllister 1933 Unknown Myrtaceae Syzygium malaccense Tiwaka Valley, New Caledonia Wood Dotte-Sarout et al. 2013 post. 650 BP Pandanaceae Pandanus tectorius Tangatatau, Mangaia Drupes Hather and Kirch 1991 ~950 BP Tangarutu, Rapa Drupes/leaf Prebble and Anderson 2012 900-320 BP Pandanus sp. Katem compound, Kosrae Parenchyma Athens et al. 1996 c.1900 BP Nikunau, Kiribati Drupes Piazza 1998 520-305 BP Kaho'olawe/Kuli'ou'ou/Anaeho'omalu Kane'ohe/Halawa/Kalahuipua'a/ Ka'ahumanu, Hawaii Drupes Allen 1984 Varied 334

Period Family Taxa Location Botanical remains Publication Dates Piperaceae Piper methysticum Nuku Hiva, Marquesas Starch Allen and Ussher 2013 550-350 BP Kaho'olawe/Kuli'ou'ou/Anaeho'omalu/ Kane'ohe/Ka'ahumanu, Hawaii Wood Allen 1984 Varied Poaceae Saccharum officinarum Kaho'olawe/Kuli'ou'ou/Kalahuipua'a, Hawaii Wood Allen 1984 Varied Rubiaceae Morinda citrifolia Katem compound, Kosrae Wood Athens et al. 1996 c.1900 BP Nikunau, Kiribati Wood Piazza 1998 520-305 BP Mauna Kea Adze Quarry, Hawaii Wood Allen 1981, 1984 ~775 BP Sapindaceae Pometia pinnata Ngofe Marsh, Vavau, Tonga Pollen Fall 2010 ca. 2500 BP

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