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THE POSSIBLE NUTRITIONAL/MEDICINAL VALUE OF SOME

TERMITE MOUNDS USED BY ABORIGINAL COMMUNITIES OF

NAUIYU NAMBIYU (DALY RIVER) AND ELLIOTT OF THE

NORTHERN TERRITORY,

WITH EMPHASIS ON MINERAL ELEMENTS.

A thesis submitted for the degree of

Master of Science

in the

University of Queensland

by

FRANCOISE L. FOTI

lngenieur Agronome (Nutrition and dietetique)

Universite Catholique de Louvain-la-Neuve (BELGIUM)

School of Chemistry and Earth Sciences

Northern Territory University

Darwin, NT

March 1994

DECLARATION

The work presented in this thesis is, to the best of my knowledge and belief, original,

except as acknowledged in the text, and that the material has not been submitted,

either in whole or in part, for a degree at this or any other university

FRANCOISE L. FOTI

Darwin, March 1994

To Uncle Emile,

My Three Children:

Nadia, Y asmin and Joshua

and Hubby Tony,

With Love •••

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(vii)

ACKNOWLEDGMENTS

A great number of people 4ave contributed to this project. Firstly, I wish to thank my

supervisor Dr David Parry who has been a constant source of guidance, constructive

criticism and reassurance and for patiently deciphering my Frenglish drafts.

Undoubtedly his support has largely contributed to the completion of this thesis.

I am grateful for the support of the Northern Territory University through the award of

the Herbalife Company scholarship which enabled me to pursue my interest in the role

of termitaria in relation to nutrition.

I wish to thank Professor David Wigston for offering me this project and making

available the use of laboratories and technical equipment.

I am also indebted to Dr Gordon Duff, Northern Territory University for his statistical

analysis rescue during the Christmas holidays and to Dr Andy Lee, Northern Territory

University for statistical advice at the beginning of the project.

Special thanks to the Daly River (Nauyiu Nambiyu) and Elliott communities and in

particular to Patricia Marrfurra Me Taggart, Molly (Yawalminy), Mercia (Wawurr) of

the Moil people in Daly River, Eileen Farrelly (adult educator of the Majellan Centre

in Daly River), Eleanor Brooks of Melville Island and Amy Lauder, Molly Dixon and

Lucy Hughes (Lababi) of Elliott for sharing their knowledge and for their support and

trust.

I am very grateful to the Aboriginal Pharmacopoeia Committee members for initial grant

and suggesting this project to Professor David Wigston; particularly Andy Barr (the

project manager) who introduced me to the communities, late Joan Chapman for her

enthusiasm, advice and encouragement and Nick Smith for sharing information and data

on Aboriginal medicine.

(viii)

A special thank you to the CSIRO Entomology Division, Canberra and especially to

Leigh Miller for the identification of termites and for sending me useful literature; to

Dr Tony Watson for giving me the opportunity to participate with his team in a field trip

and sharing his knowledge and to Dr Michael Lenz who sent me numerous photocopies

of literature articles.

I also thank the CSIRO, Division of Wildlife and Ecology, Darwin, especially Dr Gary

Cook, Andy Chapman, Dr Martin Andrew (Associate Director Roseworthy Agricultural

College) and the librarian Annette.

Thank you to Dr Barry Noller (Department of Mines and Energy) and Naseen Peerzada

(Northern Territory University) for advice on soil extraction methods and Sue Wigston

(Northern Territory Conservation Commission) and Peter Me Far lane for their assistance

in the particle size analyses.

I am grateful for the encouragement and advice received from Dr Amanda Lee of the

Menzies School of Health Research, Dr P Scheelings (Regional Director of Australian

Government Analytical Laboratories), J Leadbeater (Senior trace element analyst) and

Dr J Brand (Sydney University).

I am also indebted to Ann Alderslade and Robert Boot from the Royal Darwin Hospital

Library.

For her fantastic photographs taken in Daly River, Gaye Pascoe has to be complemented

and gratefully thanked.

Collecting termite mound samples was a laborious process made easier by the

contribution of Ann Tobin, Jan Holland, Penne Kennedy, Nicki Hanssen, Natalie Jenkins

(who also skilfully developed the black and white photographs) and in particular by

Catherine Suringa who also provided valuable assistance in the early stages of the

laboratory work.

(ix)

A special thank you to Sue Renaud (Northern Territory University) for her friendship,

support particularly at crisis time and her practical suggestions during the late stage of

the thesis.

I wish to thank all the Myilly Point lecturers, technicians, secretaries and postgraduate

students for their professional support and friendships and in particular the late Stephen

Brand for his valuable advice on Atomic Absorption Spectrophotometry, Anna Padovan

for sharing her AAS experience, Niels Munksgaard for sharing his laboratory and his

wealth of technical knowledge, Neil Smit for numerous specially brewed cups of coffee

and for letting me use his room, Dr Keith McGuinness for statistical advice, Tony

O'Grady for offering assistance and Penne Kennedy for her computer skills and constant

support.

I am very grateful to Louise Me Kenna (Director of the Harry Giese Centre), for her

support, practical advice and encouragement, and in particular strategies for balancing

responsibilities of family and study.

I am indebted to Nicki Hanssen who supported me on many occasions throughout the

entire project from field trip to proof reading part of the thesis. Her continuing

friendship is highly valued.

Throughout the years many friends have provided practical help and support, Jennifer

Fern with moral support and baby-sitting at short notice and Phil York-Barber who also

assisted with some proof reading.

Finally, I am very grateful to my three children Nadia, Yasmin and Joshua for their

generous love, patience and understanding and to my husband Tony who put up with

so many crises and doubts, who cooked so many wonderful meals and took over the

household for the last four months of the study.

(x)

ABSTRACT

This study is the first detailed investigation of the possible nutritional/medicinal value

of termite mounds as eaten by Aboriginal people. Two communities of the Northern

Territory (Nauiyu Narnbiyu, Daly River and Elliott) were chosen for detailed

investigations on the choices, usages and modes of preparation of termitaria. The

elements studied were AI, Ca, Co, Cu, Fe, K, Mg, .Mn, Na and Zn together with the

particle size analyses.

In Daly River, the Aboriginal women did not chooseAmitermes vitiosus at all sites and

preferred the mounds of Tumulitermes pastinator and Nasutitermes triodiae with the

newly built material being the most favoured. In Elliott,Amitermes vitiosus mounds

were used exclusively. In the two communities, the use of termitaria for gastric

disorders or after eating certain foods like yams, turtle or goannas could be related to

the clay content and in particular to the kaolin; while the consumption of termite mound

material during pregnancy or lactation could be associated with their elemental content,

in particular, iron and calcium.

The results of this study showed that at all sites and for all species, the mounds selected

by the Aboriginal people had a higher percentage of clay than the adjacent top soil (0-

IOcm) and the species most favoured by the Daly River people (Nasutitermes triodiae),

had the highest mean clay content. The average clay content of soil was 12.9 ± 4.4%

and for Amitermes vitiosus, Tumulitermes pastinator and Nasutitermes triodiae mounds

was: 16.3 ± 4.6, 20.8 ± 6.2 and 22.8 ± 4.2% respectively. Differences in clay content

occurred between mounds of the same species at most sites and between the same

species at different sites for Nasutitermes triodiae and Tumulitermes pastinator. With

respect to the Aboriginal preference for newly built as opposed to older material, no

significant differences were found. The clay in the tennitaria appeared to be largely

composed of kaolin. This is significant as kaolin has long been used for the treatment

of gastric-disorders in both traditional and modern pharmacologies.

(xi)

The concentrations of elements from the water "infusion" extract from Amitermes

vitiosus mounds, which is drunk by the Aboriginal people, were minimal compared to

the human needs but, nevertheless, they could contribute to the global intakes, especially

calcium. On the other hand, the finer fraction which includes clay, preferentially

selected in the infusion, could be beneficial against gastro-intestinal disorders.

In general, the termitaria had higher concentrations of elements than the adjacent top-soil

and the differences between mounds and soils were more pronounced in the bioavailable

tests. This could be a reflection of the termite by-products (of organic origin) added to

the mounds.

In the bioavailable analyses, the differences in concentrations of elements cannot always

explain the Aboriginal preference of one species at the exclusion of another at the same

site or a particular species (Amitermes vitiosus) at one site but not at another. Between

different age materials, with the exception of the soluble iron inNasutitermes triodiae

mounds, no significant differences in concentration were observed. The 44 % increase

in soluble iron in the newly built parts of mounds in the pepsin-HCl extracts could be

of importance in relation to the Aboriginal use of termitaria during pregnancy.

Undoubtedly one of the most remarkable aspects of termitaria are their high

concentrations of elements. However, with the exception of calcium which had a high

percentage recovery (82 %) between "total" and bioavailable analyses, only a fraction

of the "total" concentrations of elements present in the termitaria was potentially

available. In relation to human needs, the daily average quantity of termite mound

consumption, estimated at 30-60 g per day, can only provide a small portion of the RDis

for Ca, Cu, Fe, Mg and Mn. Fifty grams of termitaria would provide less than 5 % of

the RDis for Ca, Cu and Mg. The relatively low concentrations of Na and K in

termitaria would not provide a significant contribution towards salt replacement in the

diet. The "total" aluminium concentrations were high (2676-7745 mg/lOOg) in the

termitaria, however, only 0.37 to 13.9 mg/lOOg were present after the in vitro

bioavailable analyses.

(xii)

The bioavailability study showed that a maximum of 0.25 mg/1 OOg of iron from termite

mounds could be bioavailable to adult males. This represents less than 0.1 % of the

11total11 iron present in the termitaria studied. As the daily average loss for men is only

I mg/day, 50 g of termitaria could possibly contribute significantly to the daily

requirements. Since the bioavailability of ifon is influenced by a number of factors,

including the diet, further study will be necessary to assess more precisely the iron

bioavailability from termitaria for Aboriginal people.

(xiii)

TABLE OF CONTENTS

Page

Acknowledgments Vll

Abstract X

Table of Contents Xlll

List of Tables xxii

List of Figures xxxii

List of Plates xxxvi

, Abbreviations xxxviii

1 INTRODUCTION 1

1.1 Medicinal/Nutritional Usages of Termitaria 1

1.1.1 In Australia I

1.1.1.1 Modes of Preparation 5

1.1.2 In Two NT Aboriginal Communities 7

1.1.2.1 Nauiyu Nambiyu (Daly River) 7

1.1.2.2 Elliott I I

1.1.3 Other Countries 12

1.1.3.1 Usages 13

1.1.4 Possible Therapeutic Activities 14

1.1.5 Possible Complications 16

1.1.6 Geophagy and its Relation to Mineral

Deficiency 17

1.2 Nutritional Aspects of Selected Elements and

Recommended Dietary Intakes (RD!s) 20

(xiv)

Page

1.3 Some Aspect of Traditional Aboriginal Health

Concept and Background 25

1.3.1 Aboriginal Health Concept 25

1.3.2 Aboriginal Mineral Nutritional Background 25

1.4 Termitaria Biological Background 27

1.4.1 Taxonomy and General Biology of Tennites 27

1.4.1.1 Feeding Habits 32

1.4.2 Nests 33

1.4.2.1 Mound Construction 35

1.4.2.2 Age of the Termitaria 35

1.4.2.3 Termite Nest Material and Fabrics 36

1.4.2.4 Chemical Analyses 41

1.4.2.5 Agricultural Uses of Termitaria 68

1.5 Aims of this Project . 69

2 MATERIAL AND METHODS 71

2.1 Collection of Terrnitaria 71

2.1.2 Method of Collection 71

2.1.2 Site Locations 71

2.1.3 Sample Description and Summary 73

2.1.3.1 Detailed Mound Study 73

2.1.3.2 T ermitaria Sample 76

2.1.3.3 Soil Sampling 80

2.2 Sample Preparation 80

2.3 Particle-size Analysis

2.4 Acid Extractable of T erm.itaria and Soils

2.4.1 Extraction Trials

2.4.2 Acid Extraction Nitric/Perchloric acid (1 :4) Method

2.4.3 Atomic Absorption Spectrophotometer Analysis Procedures

2.4.4 Quality Assurance and Quality Control for the Analyses

2.5 Infusion (Hot Water) Extractable Minerals

2.5.1 Extraction Process

2.5.2 Analysis Procedures

2.6 Bioavailability of Fe in Termitaria

2.6.1 Bioavailability Extraction Trials

2.6.2 Extraction Procedure


2.6.3.1 Total Concentrations

2.6.3.2 Analysis of Fe(Il)

2.6.4 Quality Assurance and Quality Control

3 RESULTS

3.1 Site and Mound Characterisations

3 .1.1 Site Characterisations

3.1.2 Termite Species and Mound Characterisations

3.1.2.1 Nasutitermes triodiae (Froggatt)

(xv)

Page

81

81

82

82

83

85

85

85

86

87

87

88

89

89

90

90

91

91

91

91

95

(xvi)

3.2

3.1.2.2

3.1.2.3

3.1.2.4

Tumulitermes pastinator (Hill)

Amitermes vitiosus Hill

Tumulitermes hastilis (Froggatt)

Acid Extractable (Perchloric/Nitric Acids) Selected Elements from

Termite Mounds and Soils Together with Particle Size

3.2.1

3.2.2

3.2.3

3.2.4

Quality Assurance and Quality Control

Overview, General Correlation

Detailed Mound Study

3.2.3.1

3.2.3.2

3.2.3.3

Hypotheses

3.2.4.1

3.2.4.2

3.2.4.3

3.2.4.4

3.2.4.5

Amitermes vitiosus (Elliott, Site 5)

Tumulitermes pastinator (Daly River, Site 3)

Nasutitermes triodiqe (Daly River, Site 3)

Hmothesis 1: The New Material of Nasutitermes

triodiae Mounds Contains a Higher Element

Content, in Particular Iron and Calcium, and has

a Higher Clay and Silt Content than the Older

Part of the Mounds

Hypothesis 2: There is No Difference Between

Samples Taken from Different Positions of

T ermitaria for Selected Elements and

Particle Size Content

Hypothesis 3: There are No Significant Selected

Elements and Particle Size Differences Between

Mounds of Different Sizes

H.xnothesis 4: There are Differences Between

Mounds of the Same Species at the Same Site

Hypothesis 5: There are Differences in Selected

Elements and Particle Sizes Between Termitaria

of Different Species at the Same Site

Page

97

97

99

101

101

103

106

106

109

113

117

117

121

127

127

130

3.2.4.6

3.2.4.7

Hypothesis 6: There are Differences in Element

and Particle Size Content for Same Species

Mounds at Different Sites

Hypothesis 7: There are Differences in Elements

and Particle Sizes Between Different Species at

Different Sites

3.3 Hot Water ("Infusion") Extractable Selected Elements from Amitermes

vitiosus Mounds (Elliott, Site 5)

3.3.1 Comparison of Hot Water ("Infusion") Element Extracts and

Perchloric/Nitric Acid Extracts

3.3 .2 Position Effects on Selected Elements

3.4 Soluble Iron, lonisable Iron and Selected Element Concentrations of

(xvii)

Page

140

145

152

152

153

Termitaria and Soils Following Pepsin-Hydrochloric Acid Incubation 154

3.4.1 Pepsin Concentration Effects on Soluble Iron, lonisable Iron

and Selected Element content Following Pepsin-Hydrochloric

Acid Incubation 154

3.4.2 Quality Assurance 157

3.4.3 Soluble Iron, Ionisable Iron and Selected Element Composition

of Pepsin-Hydrochloric Acid (pH 1.35) Extracts and pH 7.5

Filtrates 159

3 .. 4.3.1 Soluble Iron, lonisable Iron and Selected

Elements Comparisons of Pepsin-Hydrochloric

Acid pH 1.35 Extracts, pH 7.5 Filtrates

and Perchloric/Nitric Acid Extracts 159

3.4.2.2 Age Effects on Selected Elements (Depth�O) 171

(xviii)

4

4.1

4.2

4.2.1

3.4.3.3 General Overview of the different species studied

at different sites with Relation to the Adjacent

soil (0-!0cm)

DISCUSSION

Introduction

Acid Extractable (Perchloric/Nitric Acid) Selected Element

Concentrations Together With Particle Size

Quality Control and Quality Assurance

4.2.2

4.2.3

4.2.4

4.2.5

4.2.6

4.2.7

4.2.8

Overview, General Correlation

Influence of Age of Mound Material on Selected Element

Concentrations and Particle Size

Influence of Depth in Mound on Selected Element and

Particle Size

Influence of Position in Mound on Selected Element

Concentrations and Particle Size

Influence of Mound Size on Selected Element Concentrations

and Particle Size

Comparison of Mounds of the Same Species at the Same Site

Comparison Between Different Species Mound Composition

at the Same Site

4.2.8.1

4.2.8.2

Comparison Between Tumulitermes pastinator

and Tumulitermes hastilis Mounds Composition

in Daly River Site I

Comparison Between Nasutitermes triodiae and

Tumulitermes pastinator Mounds Composition

in Daly River site 1 and Howard Springs

Page

171

175

175

175

175

177

179

181

182

184

184

185

186

186

4.2.8.3 Comparison Between Amitermes vitiosus and

Nasutitermes triodiae Mound Composition in

(xix)

Page

Daly River Site 4 187

4.2.9 Comparison Between Mound Composition of the Same

Species at Different Sites 188

4.2.9.1 Comparison Between Mound Composition of

Amitermes vitiosus at Different Sites 188

4.2.9.2 Comparison Between Mound Composition of

Tumulitermes pastinator and Nasutitermes

triodiae at Different Sites

4.2.10 General Overview: Influence of Soil Composition on

Composition of Mounds of Different Species at Different

Sites

4.3 Hot Water ("Infusion11) Extractable Selected Element Concentrations

from Amitermes vitiosus Mounds (Elliott, Site 5)


190

191

194

Termitaria and Soils Following Pepsin-Hydrochloric Acid Incubation 195

4.4 . I Quality Assurance 196

4.4.2 Selected Element Comparisons of Pepsin-HCI (pH 1.35)

Extracts, pH 7.5 Filtrates and Perchloric/Nitric Acid Extracts 197

4.4.2.1 Influence of Age of Mound Material on Selected

Element Concentrations and Particle Size,

Depth�o) 198

4.4.2.2

4.4.2.3

Comparison Between Different Species Mound

Composition at the Same Site

Comparison Between Mound Composition of the

Same Species at Different Sites

199

201

(xx)

4.4.2.4 General Overview: Comparison Between Mound

Composition of Different Species Studied at

Different Sites and their Relation to the Adjacent

Page

Soil (0-!0cm) 202

4.4.2.5

4.4.2.6

5 CONCLUSIONS

REFERENCES

APPENDICES

'Bioavailable' Composition of Different Mounds

at Different Sites in Relation to Human Needs

and Foods

Soluble Iron and lonisable Iron in Relation to the

Human Needs

I Detailed Results of Particle Size Content (%) ofAmitermes

vitiosus, Tumulitermes pastinator, Nasutitermes triodiae and

Tumulitermes hastilis Termite Mounds Sampled at Site 1-7

II Detailed Results of Selected Elemental Composition (mg/lOOg) of

Amitermes vitiosus Termite Mounds Sampled at Elliott (site 5)

Following Hot Water "Infusion"

III Detailed Results of Selected Elemental Composition (mg/IOOg) of

Amitermes vitiosus, Tumulitermes pastinator, Nasutitermes triodiae

and Tumulitermes hastilis Termite Mounds Sampled at Site 1-7,

Following Perchloric/Nitric Acid (4:1) Extraction

204

206

209

215

237

237

250

251

IV Detailed Results of Selected Elemental Composition (mg/1 OOg) of

Amitermes vitiosus, Tumu!itermes pastinator, Nasutitermes triodiae

and Tumulitermes hasti!is Termite Mounds Sampled at Site 1-6,

(xxi)

Page

Following Pepsin-HCl Extraction (pH 1.35) 265

V Detailed Results of Selected Elemental Composition (mg/1 OOg) of

Amitermes vitiosus, Tumulitermes pastinator, Nasutitermes triodiae

and Tumulitermes hastilis Termite Mounds Sampled at Site 1-6,

Following Pepsin-HCl Extraction (pH 1.35) and Neutralisation (pH 7.5) 270

VI Graphic Representations of the Soil - Mound Effects (mean ± SE) on

Element Concentrations (mg/l OOg) Following the/1.1 Vitro Test on

Tennitaria of Amitermes vitiosus, Tumulitermes pastinator,

Tumulitermes hastilis, Nasutitermes triodiae, and Soil (0-lOcm),

Sampled at Sites 1-6, following Pepsin-HCl (pH 1.35) Extractions 275

VII Graphic Representations of the Soil - Mound Effects (mean ± SE) on

Element Concentrations (mg/1 OOg) Following the In Vitro Test on

Termitaria of Amitermes vitiosus, Tumulitermes pastinator,

Tumulitermes hastilis, Nasutitermes triodiae, and Soil (0-1 Ocm),

Sampled at Sites 1-6, following Pepsin-HCI (pH 1.35) Extractions

and neutralisation (pH 7.5) 279

(xxii)

LIST OF TABLES

Page

Table 1.1 Summary of reported usages of termitaria 3

Table 1.2 Nutritional aspects of selected elements: adult body content,

physiological functions, deficiency symptoms, daily losses,

% absorption from food, recommended dietary intakes

(RDis) and sources 24

Table 1.3 Australian termite mound and adjacent soil physical

properties 44

Table 1.4 Means and standard deviations for a number of species of

Australian termite mounds {and corresponding soils)

chemical analySes 49

Table 1.5 Termite mound and soil chemical data I (Lee and Wood,

197lb) 51

Table 1.6 Coptotermes acinaciformis (Australian mound and soil

chemical analyses) 55

Table 1.7 Amitermes vitiosus (Australian mound and soil chemical

analyses) 56

Table 1.8 Nasutitermes triodiae {Australian mound and soil chemical

analyses) 60

Table 1.9 Tumulitermes pastinator (Australian mound and soil chemical

analyses) 61

Table 1.10 Tumulitermes hastilis (Australian mound and soil chemical

(xxiii)

Page

analyses) 62

Table 2.1 Site locations 72

Table 2.2 Physical characteristics of 3 termitaria selected for more

detail sampling: nasutitermes triodiae from Daly River

(site 3), Tumulitermes pastinator from Daly River (site 3)

and Amitermes vitiosus from Elliott (site 5)

Table 2.3 Detail mound sample summary for 3 termitaria:

nasutitermes triodiae from Daly River (site 3},

Tumulitermes pastinator from Daly River (site 3) and

Amitermes vitiosus from Elliott site 5)

Table 2.4 Soil samples collected at 0-1 Ocm depth in Daly River

(site 3) and in Elliott (site 5)

Table 2.5 Amitermes vitiosus termitaria sample summary

Table 2.6 Tumulitermes pastinator termitaria sample summary

Table 2.7 Nasutitermes triodiae termitaria sample summary

Table 2.8 Tumulitermes hastilis sample summary

Table 2.9 AAS Instrument Parameters

Table 2.10 AAS Instrument Parameters for Analysis of Hot Water

74

75

75

77

78

79

80

84

Digest 87

(xxiv)

Page

Table 2.11 Bioavailable test sample list 88

Table 2.12 ICP-AES instrument parameters (pepsin-HCI extraction)

(pH 135 and pH 7.5) 89

Table 3.1 Quality control of selected elemental composition of reference

material (BCSS-1, MESS-I and IAEA SOILS), following

perchloric/nitric acid ( 4: I) extraction (mg/IOOg) 100

Table 3.2 Quality control of selected elemental composition of internal

reference termitaria material (Av44D4, Av22E, Tp23Dl and

Nt24D4), following perchloric/nitric acid (4:1) extraction

(mg/IOOg) 102

Table 3.3 Quality control-of selected elemental composition of internal

reference soil material (OlE, 25D4 and 29H), following

perchloric/nitric acid (4:1) extraction (mg/IOOg) 104

Table 3.4 Pearson correlation (PC) matrix and probabilities (P) of

selected elements and particle sizes of 87 termite mounds

(n�I89) of all the species and sites studied (depth�!) 105

Table 3.5 Amitermes vitiosus mound detailed study (Elliott, site 5):

depth effects on selected elements (mg/IOOg) and particle

sizes (%) (mean ± standard deviation) together with ANOVA

probability of differences (P) between depths 107

Table 3.6 Amitermes vitiosus mound detailed study (Elliott, site 5):

position effects on selected elements (mg/IOOg) and particle

sizes (%) (meao ± standard deviation) together with ANOVA

probability of differences (P) between positions 109

Table 3.7

Table 3.8

Table 3.9

Table 3.10

Tumulitermes pastinator moWld detailed study: age effects

on selected elements (mg/IOOg) and particle sizes(%)

(mean ± standard deviation) together with ANOV A

probability of differences (P) between ages. (Depth=O)

Tumulitermes pastinator mound (Daly River, site 3) detailed

study: depth and position effects on selected elements

(mg/IOOg) and particle sizes(%) (mean± standard deviation)

together with ANOV A probability of differences (P) between

depths and positions (at depth=!)

Nasutitermes triodiae mound detailed study: age effects .

on selected elements (mg!IOOg) and particle sizes(%)

(mean ± standard deviation) together with ANOV A

probability of differences (P) between ages. (Depth=O)

Nasutitermes triodiae mound (Daly River site 3) detailed

study: depth and position effects on selected elements

(mg!IOOg) and particle sizes(%) (mean± standard deviation)

together with ANOV A probability of differences (P) between

depths and positions (at depth=!)

Table 3.11 Age effects on selected elements (mg/IOOg) and particle

sizes(%) (mean± standard deviation) in 5 Nasutitermes

triodioe mounds (Daly River, site 4). ANOV A probability

of differences (P) between ages. (Depth=O)

(xxv)

Page

Ill

112

115

116

119

(xxvi)

Table 3.12 Age (new, old) and position (top, ntiddle, bottom) effects

Table 3.13

Table 3.14

Table 3.15

Table 3.16

Table 3.17

on selected elements (mg/lOOg) and particle sizes(%)

(mean ± standard deviation) in 5 Nasutitermes triodiae

mounds sampled at site 4. ANOV A probability of differences

(P) between ages. (Depth=O)

Matrix of Pairwise Comparison Probabilities (P) (Tukey test)

between different positions of material (T =top, M=middle,

B=bottom) and age (o=old, n=new) for aluminium content

in Nasutitermes triodiae mounds sampled at site 4

Position effects on selected elements (mg/lOOg) and particle

sizes(%) (mean± standard deviation) in Amitermes vitiosus

mounds sampled at sites 2, 4 and 5. ANOV A probability of

differences (P) between positions

Position effects on selected elements (mg/1 OOg) and particle

sizes(%) (mean ± standard deviation) in Tumulitermes

pastinator mounds sampled at sites 1 and 6. ANOV A

probability of differences (P) between positions

Position effects on selected elements (mg/lOOg) and particle

sizes(%) (mean± standard deviation) in Nasutitermes

triodiae mounds sampled at sites 4 and 6. ANOV A

probability of differences (P) between positions

Pearson correlation (PC) and probability (P) matrix of mound

size (height + circumference) with selected elements and

particle sizes of three species at different sites

Page

120

121

122

125

126

129

Table 3.18

Table 3.19

Table 3.20

Table 3.21

Table 3.22

Table 3.23

Probability differences (ANOV A) [P] between mounds of

each species (Av, Tp, Nt and Th) per site for selected

elements and particle sizes

Selected elements (mg/lOOg) and particle sizes(%)

(mean ± standard deviation) of Tumulitermes pastinator and

Tumulitermes hastilis mounds sampled at site I .

(xxvii)

Page

130

Probability of differences (P) between the two species mounds 136



Nasutitermes triodiae mounds sampled at site 3.



(mean ± standard deviation) of Amitermes vitiosus and


Probability of differences (P) between species mounds




138



(mean ± standard deviation) of Amitermes vitiosus mounds

per site (2, 4 and 5) and for all sites (2+4+5) together with

the probability of differences (P) between sites 141

(xxviii)

Table 3.24 Selected elements (mg/lOOg) and particle sizes(%)

(mean ± standard deviation) of Tumu/itermes pastinator mounds

per site (1, 3 and 6) and for all sites (I+ 3+6) together with

the probability of differences (P) between sites

Table 3.25 Selected elements (mg/IOOg) and particle sizes(%)

(mean ± standard deviation) of Nasutitermes triodiae mound

samples per site (3, 4, 6 and 7) and for all sites (3+4+6+7)

Page

142

together with the probability of differences (P) between sites 144


(minimum, maximum and mean ± standard deviation) of

Amitermes vitiosus, Tumulitermes pastinator and Nasutitermes

triodiae mounds sampled at sites 1 to 7, together with the

probability of differences (P: Av-Tp-Nt) between the three

species and the pairwise comparison probabilities between

species (P: Av-Tp, P: Av-Nt and P: Tp-Nt) 146


(mean ± standard deviation) of soil samples (0-!0cm) collected

at all the site studied: 1 to 7


(minimum, maximum and mean ± standard deviation) of

soil (0-!0cm) sampled at sites I to 7, together with the

probability (P) of differences between soils

Table 3.29 Probability of differences (P) between soil and termite mound

by location and species together with the differences (%)

between the soil and the termite mound

147

148

149

(xxix)

Page

Table 3.30 Selected element concentrations (mean± standard deviation in

mg/IDOg) following hot water ("infusion") and acid

(perchloric/nitric acid) extractions of eleven Amitermes vitiosus

mounds (Elliott, Site 5) and soils, together with the percentage

recovery between extractions !53

Table 3.31 Position effects on selected elements (mean ± standard

deviation in mg/lOOg) in eleven Amitermes vitiosus mounds

(Elliott, Site 5) following two types of extraction: hot water

("infusion") extraction and perchloric/nitric acid extraction.

ANOV A probability of differences (P) between positions !54

Table 3.32 Selected element composition (mg/lOOg) of tennitaria reference

material (Nt26D4), following O.IN HCl acid (pH 1.35)

extraction with different pepsin percentage v/w (0%, 0.1%

and 0.5%), together with the probability (P) of differences

between pepsin concentrations !55

Table 3.33 Internal quality control of selected elemental composition

(mg/IOOg) of pepsin-HCl acid (pH 1.35) extracts and pH 7.5

filtrates of terntitaria sample (Nt26D4) 158

Table 3.34 Soluble iron and ionisable iron content (mean ± standard

deviation in mg/1 OOg) of termitaria and soils, in perchloric/nitric

extracts, pepsin-hydrochloric (pH 1.35) extracts and pH 7.5

filtrates together with the percentage recovery between ionisable

iron and soluble iron in pepsin-HCl extracts and pH 7.5

filtrates. Depth= 1, n=3 unless indicated 160

(xxx)

Table 3.35 Comparison of selected element concentration (mean± standard

deviation in mg/lOOg) of termite mounds (Tumulitermes

pastinator and Tumulitermes hastilis) and soils (0-lOcm),

sampled from Daly River (site I) io: A- pepsin-HCI acid

(pH 1.35) extracts; B- pH 7.5 filtrates and C- perchloric/nitric

acid (4:1) extracts, together with the %recovery between

treatments


deviation in mg/1 OOg) of termite mounds (Amitermes vitiosus)

and soils (0-1 Ocm), sampled from Daly River (site 2) and

Elliott (site 5) in: A- pepsio-HCI acid (pH 1.35) extracts;

B- pH 7.50 filtrates and C- perchloric/nitric acid (4:1) extracts,

together with the % recovery between treatments

Table 3.37 Comparison of selected element concentration (mean ±standard


pastinator and Nasutitermes triodiae) and soils (0-lOcm),

sampled from Daly River (site 3) io: A- pepsin-HCI acid


acid (4:1) extracts together with the% recovery between

treatments


deviation in mg/1 DOg) of termite mounds (Nasutitermes

triodiae) and soils (0-!0cm), samples from Daly River (site 4),

depth�O (new/old material), io: A- pepsio-hydrochloric acid


acid (4:1) extracts together with the% recovery between

treatments. Probabilities (P) of differences between ages

Page

163

164

165

(new/old) 166


deviation in mg/1 DOg) of termite mounds (Amitermes vitiosus

and Nasutitermes triodiae) and soils (0-IOcm), samples from

Daly River (site 4), in: A- pepsin-HCl acid (pH 1.35) extracts;

B- pH 7.50 filtrates and C- perchloric/nitric acid (4:1) extracts,

together with the % recovery between treatments

Table 3.40 Comparison of selected element concentration (mean ± standard


pastinator and Nasutitermes triodiae) and soils (0-IOcm),

samples from Howard Springs (site 6), in: A- pepsin-HCl

Table 3.41

Table 3.42

Table 3.43

Table 4.1

(pH 1.35) extracts; B- pH 7.50 filtrate and C- perchloric/nitric

acid (4:1) extracts, together with the % recoVery between

treatments

Selected Elements (mg!IOOg) (Minimum, Maximum and

Mean ± standard Deviation) of Amitermes vitiosus,

Tumulitermes pastinator and Nasutitermes triodiae mounds

and soils (0-!0cm) sampled at Sites I to 6, Following

Pepsin-HC! Incubation (pH 1.35)

Selected Elements Content (mg/IOOg) (Minimum, Maximum

and Mean ± Standard Deviation) of Amitermes vitiosus,

Tumulitermes pastinator and Nasutitermes triodiae mounds

and soils (0-!0cm) Sampled at Sites I to 6, in pH 7.5 Filtrates

Selected Element Differences(%) Between Soil and Termitaria

in Pepsin-HCl Acid Extracts (pH 1.35) and pH 7.5 Filtrates

Composition of Selected Australian Aboriginal Bushfoods

and Western foods in mg per lOOg Edible Portion

(xxxi)

Page

167

168

172

173

174

205

(xxxii)

Figure 1.1

Figure 1.2

Figure 1.3

Figure 1.4

Figure 1.5

Figure 1.6

Figure 1.7

Figure 1.8

Figure 1.9

LIST OF FIGURES

Location of reported usages of termitaria throughout the

Northern Territory

Classification of lsoptera

Termite families

Castes of Coptotermes acinaciformis, Rhinotermitidae:

A: winged reproductive or alate

B: worker

C: soldier

De-alate reproductive (dropped wings) revealing severed

wing butts

Queen termite with a distended abdomen (physogastric queen)

Soldier types:

A: mandibulate soldier of Heterotermes ferox,

Rhinotermitidae

Page

2

28

29

30

30

31

B: nasute soldier of Nasutitermes exitiosus, Termitidae 31

Digestive tube system in worker termites

Mound nest:

A: Mound nest of Coptotermes lacteus

B: Mound nest of Coptotermes lacteus showing

internal stnucture

32

Figure 3.1

Figure 3.2

Figure 3.3

Figure 3.4

Figure 3.5

Figure 3.6

Figure 3.7

Figure 3.8

(xxxiii)

Page

Dorsal view of nasute head: A: Nasutitermes sp and

B: Tumulitermes sp and mandibulate head: C: Amitermes sp 93

Size comparison (height + circumference in em) of Amitermes

vitiosus (Av), Tumulitermes pastinator (Tp), Nasutitermes triodiae

(Nt) and Tumulitermes hastilis (Th) mounds at different sites 95

Position effects (mean ± SE) on calcium, magnesium and zinc

(mg/1 OOg) in Amitermes vitiosus mound sampled in Elliott

(site 5) at depths I and 2

Depth effects (mean ± SE) on calcium, potassium and

manganese (mg/1 OOg) in Tumulitermes pastinator mound

sampled in Daly river (site 3)

Depth effects (mean ± SE) on silt and coarse sand (%) in

108

110

Tumulitermespastinator mound sampled in Daly River (site 3) 110

Depth effects (mean ± SE) on iron and sodium (mg/1 OOg)

together with coarse sand (%) inNasutitermes triodiae mound

sampled in Daly river (site 3)

Position effects (mean± SE) on calcium (mg/lOOg) and coarse

sand (%) in Nasutitermes triodiae mound sampled in Daly

114

River (site 3) 114

Age effects (mean ± SE) on aluminium, iron, potassium and

copper (mg/1 OOg) in Nasutitermes triodiae mounds sampled

in Daly River (site 4) 118

(xxxiv)

Figure 3.9 Age effects (mean ± SE) on aluminium and iron (mg/1 OOg),

at three positions (top, middle, bottom), inNasutitermes

triodiae mounds sampled in Daly River (site 4)

Figure 3.10 Position effects (mean ± SE) on calcium, magnesium and

manganese (mg/1 OOg) in Amitermes vitiosus mounds

sampled in Daly River (site 4)

Figure 3.11 Position effects (mean ± SE) on particle size ( (%) in

Page

118

124

Amitermes vitiosus mounds sampled in Daly River (site 4) 124

Figure 3.12 Mound effects (mean± SE) on calcium and iron (mg/IOOg)

in Nasutitermes triodiae mounds sampled in Daly River

(site 4)

Figure 3.13 Selected elements (mg/IOOg) and particle size (%) (mean± SE)

of Amitermes vitiosus (Av), Tumulitermes pastinator (Tp),

Nasutitermes triodiae (Nt) and Tumulitermes hastilis (Th):

128

A Aluminium and calcium concentrations (mg/1 OOg) 131

B Cobalt, Copper, Iron and potassium

concentrations (mg/IOOg)

C Magnesium, manganese, sodium and zinc

concentrations (mg/1 OOg)

D Clay, silt, fine sand and coarse sand content (%)

Figure 3.14 Soil-mound effects (mean± SE) on calcium, potassium and

sodium (mg/100g) in mounds of different species and soils

(0-1 Ocm) sampled at different locations

132

133

134

ISO

\

Figure 3.15 Soil-mound effects (mean ± SE) on clay and coarse sand (%)

in mounds of different species and soils (0-lOcm) sampled at

different locations

Figure 3.16 Pepsin concentration effects on aluminium, soluble iron and

ionisable iron (mean± SE in mg/lOOg) inNasutitermes

triodiae mound (Nt26D4, Daly River, Site 4) following

Pepsin-HC1 pH 1.35 extraction (n�9)

Figure 3.17 Pepsin concentration effects on aluminium, soluble iron and

ionisable iron (mean ± SE in mg/lOOg) inNasutitermes

triodiae mound (Nt26D4) in pH 7.5 filtrates (n�9)

Figure 3.18 Selected element percentage variations between

perchloric/nitric acid extracts, pepsin-HCI acid pH 1.35

extracts and pH 7. 5 filtrates of termitaria (A: Amitermes

vitiosus; B: Nasutitermes triodiae and C: soil , from Daly

River site 4)

Figure 3.19 Selected elements (calcium, potassium, magnesium and

manganese) percentage recovery in pH 7.5 filtrates of

Nasutitermes triodiae mounds, Tumulitermes pastinator

mounds and soils at site 3

(xxxv)

Page

151

156

157

162

170

(xxxvi)

Plate 1

Plate 2

Plate 3

Plate 4

Plate 5

Plate 6

Plate 7

Plate 8

LIST OF PLATES

Nathan (Culmungu) and Aaron (Kingaruo) Me Taggart

inspecting an Amitermes vitiosus mound in Daly River

Mercia (Wawurr) of the Moil people at Nauiyu Narnbiyu

(eating some Nasutitermes triodiae tennitaria) accompanied by

Molly Yawalminy and Malcolm Kurruwul

Typical Nasutitermes triodiae mounds in Daly River

Mercia (Wawurr), Molly (Yawalminy) and Patricia Marrfura

Me Taggart of Daly River showing the newly built material,

that they favoured, at the base of aNasutitermes triodiae

mound

A sample of Nasutitermes triodiae mound ready to be taken

home (Daly River)

Sample of termitaria quantity Mercia (Wawurr) and Molly

(Yawalminy) used to take 2 or 3 times a day during their

pregnancy (Daly River)

Lucy Hughes (Lababi) from Elliott, crushing someAmitermes

vitiosus mounds during the preparation of the "infusion"

In Elliott, Lucy Hughes demonstrates the poultice while Amy

Lauder is getting ready to drink the 11infusion"

Page

VI

4

6

6

8

8

10

10

Plate 9

Plate 10

Plate II

Plate 12

Plate 13

Plate 14

Plate 15

Plate 16

Typical site with Nasutitermes triodiae (right) and Amitermes

vitiosus (left) mounds in Daly River

Nasutitermes triodiae mound (3 m high), Daly River site 3

Vertical section of Nasutitermes triodiae mound at site 3,

showing compact basal portion-galleries and nursery (middle

part of the mound, ground level)

(xxxvii)

Page

92

94

94

Tumulitermes pastinator mound (70 em high), Daly River, site 3 96

Vertical section of Tumulitermes pastinatormound at site 3,

showing alveolar type of structure and the nursery (N)

Amitermes vitiosus mound (50 em high), Elliott site 5

Vertical section of Amitermes vitiosus mound in Elliott,

showing the concrete hard structure

Tumulitermes hastilis mound (55 em high), Daly River site 1

96

98

98

99

(xxxviii)

AAS:

ANOVA:

Av:

CEC:

cucum:

em:

diam:

Ext. Lab:

g:

ICP-MS:

ICP-OES:

Kg:

L:

m:

me:

mg:

mm:

run:

Nt:

OSS:

pmw:

ppm:

RDis:

SD:

SE:

Th: Tp:

XRF:

ABBREVIATIONS

atomic absorption spectrophotometer

analysis of variance

Amitermes vitiosus

cation exchange capacity

circumference

centimetre

diametre

external laboratory

gram

inductively coupled plasma - mass spectrometer

inductively coupled plasma - optical emission spectrophotometer

kilogram

litre

metre

milliequivalent

milligram

millimetre

nanometre

Nasutitermes triodiae

office of the supervising scientists

post-menopausal women

parts per million

recommended daily intakes

standard deviation

standard error

Tumulitermes hastilis

Tumulitermes pastinator

x- ray fluorescence

CHAPTER ONE

INTRODUCTION

1 INTRODUCTION

1

"Most nineteenth-century colonists looked with horror at the eating habits of Aborigines,

and had little insight into the depth of knowledge and experience of their hosts. "82

1.1 Medicinal/Nutritional Usages of Termitaria

This study is a part of the Aboriginal Pharmacopoeia Project which aims to study the

therapeutic use of Aboriginal plants and other natural products in Northern Australia.

1.1.1 In Australia

Like many other indigenous populations around the world, the Aboriginal communities

are using termite mounds (termitaria) for many different reasons: physical and

metaphysical. Although the spiritual and cultural aspect of the bush medicine cannot be

ignored. only its physical aspect has been addressed in this study as it can be studied by

modem scientific methods.

Termite mounds are known by many different names. Besides their local names, such

as: ''Ngete" in Ngankikurungkurr, "Bilaya" in Jingulu and "Bellar" in Mudburra1\ they

are popularly known as "ant"-bed, "ant"-hill and termite hill14• They are used

medicinally throughout the Northern Territory from Bathurst Island to the Central

Northern region14 (see Figure 1.1). Different Aboriginal communities are using

tennitaria for different purposes and in many different ways. It is used either internally

or externally. Their usage, as with other bush medicine, was more widespread prior to

European contact. As Magarruminya, a Mardarrpa (Yolngu) man said: "We had our own

medicine. We had doctors of our own ... Now young people are giving that away ... "160,

There are multiple use of termitaria; see Table 1.1 Their main applications are to treat

gastro-intestinal disorders17•44•43•82 or are related to pregnancy.14

2

...

Bathurst Island

�

Goulburn Island Warruwi

'L.:V> .. !

Pine Creek • •

Nauiyu Nambiyu

Groote Eyiandt 1 n:Umbakumba,

"1"""f'" Bulla

• ,

FIGURE 1 . 1

•

Lajamanu ..

Newcastle Waters

• . Elliott

0 •M 200 3�: km

Location of reported usages of termitaria throughout the Northern Territory

3

TABLE 1.1 Summary of reported usages of termitaria

Use Community (Figure 1.1) Reference

Diarrhoea Pine Creek, Maningrida 131 Lajamanu, Bulla, Nguiu, 43 Elliott, Newcastle Waters, 14 Nauiyu Nambiyu

Pregnancy W arruwi, Umbakumba, 14 Nauiyu Nambiyu

Abdominal or Goulburn Island 14 menstrual pains

Mineral deficiency Groote Eylandt 100

General illness Elliott 16 Bathurst Island 131

Lactation Elliott 14

Sores Bathurst Island 131

Allay hunger Nauiyu Nambiyu 14 Out-stations (Maningrida) 44

Cooking Elcho Island 16 Bathurst Island

Mosquito repellent Goulburn Island 131

Eastwell (1979)" mentioned that one bush group of 40 people consumed 2 kg of termite

mound during April 1973. Eastwell (1984)44 reported that "people who live in the out

stations still eat some ant-hill with some of their meals, or have some clay"44 and that

"teenagers are seen carrying clay in plastic bags to eat at the outdoor cinema1143•

Sometimes, in Daly River, when there is a flood that renders the termite mounds

inaccessible, the people would dive down to collect clay from a favoured place. Clay

and termite mounds seem interchangeable with preference for termitaria in some places.

Levitt (1980)100 wrote that clay processed by animals, such as on the termite mound, was

considered to be safer than other kinds. Where clay is eaten, it is used in much the

same way as termite mounds: to allay hunget2.17, to cure stomach aches, diarrhoea, to settle the stomach, to treat worm infestation or to "line the stomach before eating yams,

or fish which may be poisonous"11.

4

.. 0

2 11-0

"" .. c ~ 0

~

PLATE 2 Mercia (Wawurr) of lhe Moil people at \lauiyu -.;ambiyu (eating some Nasutitermes tnodioetennlto.ria) accompanaed by Molly (Y JWalminy) and Mnlcolm Kurruwul

5

Bateson and Lebroy (1978)17 reported that children ate clay because they liked the taste.

On Groote Eylandt, some women ate clay "particularly if they had a craving for fish as

they said it tasted like fish"100•

1.1.1.1 Modes of Preparation

There are many different ways in which the Aborigines use termite mounds. It seems

almost as if every family has its own "recipes" . The simplest way is to break off small

pieces of the outer mound, crumble it in the hand to a powder, then drop it into the

mouth (Nauiyu Nambiyu1\ Groote Eylandt100), (Plate 2). In other communities, a large

piece of mound (hand size) is ground finely and mixed with water, milk or tea, then

drunk.14•82 Sometimes, only the extracted liquid is drunk (Wave Hill, Wattie Creek)17.

Honey-ants (Melophorus sp.) or plants can be added to the mixture of tennitaria and

water16. In Elliott14, pieces of mounds are first burnt in the fire, until black, then crushed

and mixed with water. One part of the "infusion" is then drunk and the other part used

to bathe the skin or as a poultice.16 When used as a mosquito repellent, the inner part

of the mound is slowly burnt in the frre4.

In general, the mounds are consumed on the spot. In some places, such as Numbulwar,

the dry earth eaten was probably tennitaria brought from another area for the

construction of the airstrip17• On some occasions, for example, just prior to the flood

caused by the rains of the wet season in Daly River, a big lump of the mound can be

taken horne for further use.

The mound is not the only part eaten, Dulcie Levitt (1980i00 reported that the tennite

soil tunnels in logs are also used.

Since tennite mounds have been so widely used in Australia and that most soft-bodied

insects that were available in quantity were eaten190, it is surprising that there is no

mention of termites themselves being used as food. Likewise, despite the abundance of

tennitaria in the Amazon region, few records exist of Brazilian Indians eating termites110,

compared with Africa where they are a favourite dish164•

" 0 u .. .. 0.. <) ;-;-,� '-' " 0 ..::::

:::.

'-' 0 0 E .. » .. 0 0 0 "" e:.

6

PLATE 3

PLATE 4

Typical Nasutitemus triodiae mounds in Daly River.

Mercia (Wawurr). Molly (Yawalminy) and Patricia Marrfura Me Taggart of the Daly Rjver showing the newly built material, that they favoured.

at the base of a Nasutllem1es triodiae mound.

1.1.2 In Two NT Aboriginal Communities

7

The selection of the two communities (Nauiyu Nambiyu (Daly River) and Elliott) was

suggested by Andy Barr (the Aboriginal pharmacopoeia project manager) and the late

Mrs Joan Chapman (the pharmacopoeia team pharmacist) who had worked closely with

the two communities and had received the approval by the two communities for further

study.

1.1.2.1 Nauiyu Nambiyu (Daly River)

The first contact with the Nauiyu Nambiyu community took place in June 1988. A

meeting was organised with Patricia Marrfurra Me Taggart of the Moil people at Nauiyu

Narubiyu (coordinator at the MajelJan Centre), her mother Moliy (Yawalminy), her

·grand-mother Mercia (Wawurr), her two children: Nathan (Culmungu) and Aaron

(Kingaruo), and Eileen Farrelly (adult educator of the Majellan Centre).

The information concerning the use of termitaria was gathered after a few subsequent

visits and field trips (August-October-November 1988, December 1988 and 1989, August

1990 and 1991), often also including members of the extended family.

1.1.2.1.1 Choice of Termitaria

Mercia and her family use three different types of mounds. They belong to:

'Nasutitermes triodiae, Tumulitermes pastinator and Amitermes vitiosus. Where

Nasutitermes triodiae is present, Amitermes vitiosus is not taken. The red mounds may

be thought to be more effective16• Coptotermes acinaciformis mounds have also been

reported by the pharmacopoeia team as being used in Daly river. Out of al� the mounds,

they prefer the freshly built parts of Nasutitermes triodiae mounds (Plates 3 & 4).

11Women like the taste of it" (Mercia). Just seeing the freshly built parts give them

watery mouth.

8

PLATE 5

PLATE G

A sample of Nasutitermes triodiae mound ready to be taken home (Daly River).

Quantity of termitaria Mercia (Wawurr) and Molly (Yawalminy) used to

take 2 or 3 times a day during thdr pn:gnancy (Daly River)

9

If the mound is attacked by red ants they would not eat it because it would be too old.

The 11old ones (are) not too sweet" said Mercia. Sometimes, they take "a big mob

home" for further consumption (Plate 5). They throw away the grass and termites by

shaking the portion taken and let it "dry" in the sun before putting it into their bag.

When tennitaria is not available (eg: during the flood of the wet season) they dig for a

clay (sub-soil) close to the mission.

1.1.2.1.2 Usage

Mercia said that they used to eat it 11more before the mission, now they forget about it".

She used to eat a handful (20 to 30 g) (Plate 6) at least 2-3 times a day when she was

pregnant or had diarrhoea and used to give termite mound pieces even to young children

with diarrhoea. She uses the clay in much the same way. Molly ate termitaria material

for "upset stomach, vomiting and diarrhoea" or when she was pregnant to stop the

craving. Molly used to eat "bits and pieces all day long11 during pregnancy. She

reported that they also used to eat it when they were hungry, when they had stayed in

the bush for a long time, or after eating yam or meat (turtle, wallaby, goanna and

porcupine). She mentioned that men eat termite mounds too. Patricia reported eating

termite mound when she was a child.

1.1.2.1.3 Mode of Preparation and Consumption

Small pieces of the outer casing are broken off, crushed between the hands and placed

on the middle of the tongue where they are left to melt before being swallowed.

10

PLATE 7 Lucy Hughes (Lababi) from Elliott, crushing someAmitermes vltiosus mounds during the preparation of the "infusion".

PLATE 8 In Elliott, Lucy Hughes demonstrates the poultice while Amy Lauder is getting ready to drink the "infusion.

11

1.1.2.2 Elliott

The first contact with the community took place in September 1988. As for the Nauiyu

Nambiyu community the information was gathered after a few meetings and field trips

with the Aboriginal informers (Amy Lauder, Molly Dixon and Lucy Hughes (Lababi)).

A) Choice of Termitaria

Amitermes vitiosus mounds are the dominant mounds in the landscape and are used by

Aboriginal community for medicinal purposes. Their local name is: "Bilaya".

B) Usage

It is used for diarrhoea, upset stomach, during pregnancy or to bring up milk after birth

and to "make the baby strong".

C) Mode of Preparation and Consumption

The mound is first pushed to the ground and broken into big pieces with an axe. Then,

the pieces are placed in the fire. They remain there for 15 a2Q minutes before being

placed in a billycan, half-full with water. The pieces are crushed to mud with a piece

of stick (Plate 7) and left there to infuse for I 0 minutes . The "infusion" is mixed well

and drunk hot. The remaining mixture of muddy composition is rubbed on the back and

the front of the person (baby and lactating mother), (Plate 8).

12

1.1.3 Other Countries

Native populations throughout the tropical and sub-tropical regions of the world have

been using termite mounds for many different purposes. As previously discussed in

chapter 1.1, only the physical aspects of their use have been considered.

Other than being used for pottery68•86•164 or to build roads86•143•68, tennis courts86•143,

housesw3•86•42•15 and ovens103•42•15, termite mounds have been widely used in agriculture

around the world (see chapter 1.4.2.5) and in geochemical explorations137• Termite

mounds are also an important source of nutrition for many species of animals. In

Northern Ghana, the indigenous population use Trinervitermes geminatus mounds as a

source of chicken food157• Pullan (1974)147 reported the use of termite mounds for

nutrient supply, for example, elephants may excavate tennitaria200 and eat the earth, and

antelopes may use natural excavations of termite mounds as salt licks. In Guinea

(Conakry), tennite mound excavations containing large white nodules (most probably

calcium carbonate) are given to the cattle as a natural mineral supplement by the

nomadic breeders around th� country (Personal communication 1981). Those "earths"

were also fed to mineral deficient sows at the University farm of Macenta (Guinea)

(personal observation 1981 ). Monkeys have also been reported eating termite

mounds75•74•40•210•73• Red leaf monkeys (Presbytis rubicunda), in Sabah (Northern

Borneo), were observed eating Macrotermes mounds and chimpanzees were reported,

in Gabon and Tanzania, eating 10 to 20g of earth (most of them from tennite mounds),

up to twice daily74•

This phenomenon (the habit of eating earth) is more commonly known as geophagy.

It is a fonn of pica which is the craving and eating of non-natural, and inedible objects70,

It has been reported in the human population for many centuries45•187•47, It is reported

all over the world70: Saudi Arabia70, TurkeY3•114, Iran146•188•1S4,Js9, Algeria12, Nigeria, Togo,

Liberia and Ghana186•185 Zambia147 South Africa189 USA181•180'47•46•56•53 South America1 10 • • • • •

Haiti63, India86 and Philippines41• "Earth eating cross-cuts ethnic, social, and economic

lines. In the United States, geophagy is found among whites and blacks, children and

adults, and in both rural and urban populations. "187 Its prevalence varies greatly from

13

culture to culture66• It seems to be more prevalent in the black Americans47 and among

"poverty·stricken" populations where the nutrition is less than optirnum65 and in

women. 6s·46•147•66•47•45•147 For example: it occurs among 57% of women and 16% of

children of both sexes in the black population of rural Holmes County, Mississipi; the

average daily consumption is 50g187• And more particularly in women during pregnancy:

out of 42 women studied in Alabama who ate between 6-130g of clay daily, 38 were

pregnant46• In Ghana, although clay (from sub·soil or termitaria) is mostly eaten by

adult women (34 to 68%, with the highest percentage in more remote areas), it is also

consumed by 14% of the adult male population (in Eweland)'"'. In South Africa, clay

from a termite mound is very popular with rural black women; 44% of rural black

women experience geophagy during pregnancy as opposed to 4.4% in mixed-coloured

women and 2.2% in Indian women; while pica did occur among white pregnant women,

it was rare1119• Cavdar et al (1983)33 report that geophagy was a common finding among

Turkish children and women in villages.

In some small southern towns in the USA, clay can be purchased at commercial

outlets46• In African countries it can be bought in the local market where a particular

clay called eko or Calabar could be imported from a distant region (more than 1500 Km away)186• In Western countries, capsules of dirt and clay are being marketed in health

food stores. The origin of the material used is not always specified.

1.1.3.1 Usages

Cesare Bressa (early 1 800), cited in Mustacchi (1971)121 described that the slaves in

Louisiana started eating soil during their illnesses (known later on as wet beriberi or

vitamin Bl deficiency); most often they wanted to eat earth and some seemed to prefer

hard earth while others liked clay121• In India, Joseph (1978)86 reported that the material

of the termite mounds was often eaten by local tribes, presumably for their mineral salts

content. In Nigeria, they used clay in traditional medicinal preparations for intestinal

problems (stomach and dysenteric ailments) or problems associated with pregnancy186•

The reasons given by the pregnant women (USA) for eating clay are numerous: to

14

relieve nausea, prevent vomiting, relieve dizziness, relieve headaches and many reported

eating more clay when they were upset46• In the Amazon, termite mounds are prepared

as remedies for bronchitis, constipation, goitre, sores, boils, ulcers and other ailments110

1982• Termite mounds are also eaten when food is scarce1 10• The Uaica Indians chew the

soil of pulverised termitaria, saying that it is good for them, it helps strengthening and

building up the body; they like the taste and eat handfuls at a time110,

1.1.4 Possible Therapeutic Activities

The external uses of clay are perhaps more familiar; poultices containing clay are a

known remedy for boils, while mud baths are recommended for the treatment of

rheumatism and arthritis. The popular literature claims that the benefits of termite

mound and clay consumption are quite numerous. According to Dextreit (I 976)41 clay

can cure anaemia, absorb harmful substances (bacteria, poison), expel worms, relieves

digestive disorders including ulcers, enteritis, dysentery, constipation and diarrhoea and

many other ailments. Dextre�t (1976t1 claimed that clay can cure iron deficiency, not

because of a high mineral content, but because it contains catalysts that work in

infinitesimal doses to stimulate failing organs. The more clay is exposed to sun, air, and

rain, the more active it becomes41 •

The most obvious reason given by many authors for geophagy is because o f the potential

as nutrient sources, mainly during pregnancy. For example, in Zambia, pregnant women

use te�ite mounds probably as a source of calcium and sodium147• Barbier et a/

(1986Y2 suggested that malnutrition causing growth failure and zinc deficiency could be

the cause of geophagy12• Edwards et a/ (1959t6 noted that possibly clay-eaters eat clay

to compensate a diet that was poor in calcium and iron. This will be discussed in more

detail in section 1.1.6

In the treatment of stomach complaints, including diarrhoea, clay (from soil or

termitaria) may act as an adsorbent antidiarrhoeal and might help to alleviate digestive

disorders40• The mineralogical analyses of the eko clay (Nigeria) are strikingly similar

15

to the clay in the commercial pharmaceutical Kaopectate (kaolinic composition)m.

Kaolin is a powdered hydrated aluminium silicate freed from gritty particles171• It has

long been used for the treatment of gastric disorders (diarrhoea, dysentery) in both

traditional (China) and modem pharmacologies178• The clay absorbent power is quite

extraordinary, according to Dextreit (1976t1, 5g of clay are enough to completely

discolour 10 cm3 of a water solution containing 0.1% of methyl blue41•

An interesting case, demonstrating the absorbent power of the clay, was reported by

Halsted ( 1968)", in which a prisoner, in Germany (158 1), who was given 6g of mercuric

chloride followed by one tablet of clay ("terra sigillata in olde wine") and did not die.

Halsted (1968)" reported that it is quite possible the clay had acted as an ion-exchange

resin and that elements in the clay could have exchanged with mercury, making it

unavailable for absorption.

. Research with rats has shown that geophagy occurs when rats are made acutely ill

(poison) or are stressed or arthritic32• The sicker the rat, the more kaolin it eats. Kaolin

intake is also a behavioural index of motion sickness in rats179• Geophagy could be a

response to generalised stress, since this state may involve gastro-intestinal malaise32·m.

A study done in South Africa (where geophagy is more common among black pregnant

women), on the frequency and severity of nausea and vomiting during pregnancy

reported that it affected only 3 .8 and 3 .19% of the black women respectively, while it

affected 19.8 and 17.8% of white women respectively189

Davies and Baillie (1988)'to observing monkeys in North Borneo suggested that geophagy

may serve different functions at different times: it might help to absorb toxins, to alleviate digestive disorders or to supplement the mineral intake. This latter point was

disputed by Hladik (1977ar3 who stated that the amounts of minerals in the termite

mound sample were very small but for iron, and that the iron content of the leaves and

fruits eaten by the primates was enough to cover all their physiological requirements73•

16

There may also be other determinants such as psychological and social. For one thing,

home treatment (eg: preparation and administration of the clay-tennitaria in illness) may

strengthen family ties because care and attention are exchanged within the family group.

The patient is surrounded and taken care of by his own people who feel responsibility

for the family1s health174 but also 11Eating clay is psychologically comforting.

Masticating it is gratifying, salivary flow is increased, and the stomach is comfortably

julf'43• Likewise, the most common reason for eating clay given by males (Ghana) is

that it provides "pleasure11185

1.1.5 Possible Complications

The medical literature on geophagy is meagre and research reports are often conflicting

in their findings45• In Australia, it is not viewed as a serious medical problem by the

nursing staff of different Aboriginal communities ( eg: Maningrida, Angurugu }, although

it can cause constipation when used to excess69 According to Hausheld (1975t9 it is a

traditional practice more likely to be beneficial to health than dangerous to it69•

However, Bateson and Lebroy (1978}17 (Northern Territory) warned that eating clay or

termite mounds can be dangerous as it can cause a partial obstruction, or even

perforation, of the colon. Thomson (1984}'x1 reported that geophagy, especially if kaolin

is involved, may lead to a zinc deficiency. Eastwell (1984)44 reported that it could also

interfere with iron absorption.

Dirt eating was listed as a cause of death in 1850121, probably because it was consumed

by dying slaves in their attempt to compensate their malnutrition. Geophagy is also

associated with severe iron deficiency anaemia,s6•33•188'146,n4,154 in addition to zinc

deficiency33•66• A syndrome characterised by geophagy, iron deficiency anaemia, zinc

deficiency, growth retardation and hypogonadism has been observed in both sexes in

Iran146•154•159, and in Turkey33·m for several decades; this syndrome is probably due to the

poor nutritional status of the population but possibly increased by prolonged geophagy33•

Clay ingestion has also been associated with myositis (muscle weakness) and

hypokalemia (potassium deficiency) or hyperkalemia (excess potassiumY3•81• Halsted

17

(1970t6 noted that a case of tetanus linked to the ingestion of clay containing spores of

the tetanus bacillus has been reported66•

1.1.6 Geophagy and its Relation to Mineral Deficiency

The traditional medicinal uses of termite mounds, such as for gastro-enteric disorders

and during pregnancy would suggest that a number of elements may be of importance,

in particular minerals such as: calcium, iron, magnesium, potassium and sodium. The

older literature suggests that iron-deficiency lead to geophagy. Halsted (1968)65 reported

that although it is an attractive theory, the literature evidence for anaemia resulting in

geophagy is meagre187 and the results are often conflicting. Barbier et al (1986)12

suggest that although geophagy was probably a spontaneous effort to compensate for

malnutrition (growth failure, zinc deficiency), it was also probably the cause of iron

deficiency anaernia12• This idea is supported by Cavdar er al (1983}33 who suggested

that the zinc deficiency could have been present before geophagy started. Indeed, the

nutritional status of Turkish villagers was poor: based on wheat bread and wheat product

with very limited animal protein. It had little zinc content and the zinc could have been

bound by phytate (inositol hexaphosphate) and fibres"'. But they also noted that the

geophagy may cause both iron and zinc deficiency: by reducing the appetite for normal

food, by decreasing the iron absorption and by causing malabsorption of zinc and iron

due to irreversible changes in intestinal epithelial cells associated with prolonged

geophagy. This same view is also supported by Walker et a/ (1985)189 who suggested

that "In the case of geophagy, while the practice may contribute significant amounts of

calcium and trace elements, under certain conditions it may exacerbate an existing iron

deficiency anaemia. The extent to which iron deficiency causes geophagy, or geophagy

promotes iron deficiency is not wholly clear ... ".

Different experiments have produced different results. For example, Patterson and

Staszak (1977)141 reported maternal anaemia and reduction in the birth weight of the

neonatal rats born to kaolin (20%) fed rats while the results of kaolin fed rats receiving

iron supplement were normal. Edwards et a/ (1983ts reported that small levels of clay

18

conswnption (20%) may be beneficial as it enhances physical development in rats. The

haemoglobin values of baby rats from female rats receiving 20% and 35 % of clay

through their diet were not different from control values45•

Eastwell (1979)43 studying Aboriginal clay eaters found no difference in the haemoglobin

levels of ingesters as compared with controls. Edwards et a/ (1964)47 reported that

although the total iron and calcium content of the clay studied was respectively:

20.8mg/100g and 21.4mgil 00g, only 0.03mgil00g ofiron and 0.2 mgilOOg of calcium

were "available" in the in vitro tests. Likewise, Venneer (1971)185 found very little

"available" minerals (0.1 N HCl extraction) in Ghanian clay. But he commented that

"It is possible, ... that even such small amounts may contribute to the overall nutrition

if that element is deficient in the body and if other dietary inputs is inadequate"m.

Although the ingestion of clay may slow down the mobility in the gastro-intestinal tract

and thus promoting absorption, its consumption may also impair the absorption of certain

other nutritive elements through chemical exchange185• Talkington et a/ (1970Y80

reported that while the ingestion of sizeable amounts of clays, just prior to iron intake,

did not appreciably reduce iron absorption; a red clay containing considerable iron

proved inefficient for correcting iron-deficiency anaemia while the admission of a

smaller quantity of ferrous sulfate did. They commented on the fact that iron absorption

in the same individual can vary appreciably, at different times, unrelated to the agent

being tested. They suggested that the degree of iron absorption reduction caused by clay

could vary widely, apparently depending upon the clay.

In their study on the effect of clays of different origins on 59FeS04 absorption, Minnie.

et a/ (1968)u4 reported that Turkish clay and soil reduced the amount of 59FeS04

absorbed from the intestinal tract while three other clays obtained in the United States

were less effective. The last clay tested was from New Mexico and it had no effect

upon iron absorption. Interestingly, a dose of MgO completely blocked radioiron

absorption in four of five subjects and reduced absorption significantly in the fifth.

Turkish clay and soil also removed iron from solution better than did the naturally acid

clays. The mechanism which most likely accounts for the observed effect is the cation

exchange capacity and the base saturation value (which correlate with the pH114). In the

19

study by Minnie et a/ (1968)1" the clay differed in effect depending upon their cation

exchange capacity (CEC). Turkish clay having a high CEC was more effective in

blocking iron absorption than were three other clays with lower CEC values. The iron

is exchanged for Ca, Mg, Mn, Na, K, and H ions with the formation of non-absorbable

iron compounds. They commented that the effect of clay and soil on iron absorption

may not be the only factor in the production of anaemia in geophagy, but it could be

contributory. Nutritional and parasitic (worms) factors may contribute to anaemia as

wel1114•

Halsted (1968)65 suggested that as geophagy leads to iron deficiency, it may also prevent

absorption of potassium and mercury and possibly zinc, as they have demonstrated a

high cation-exchange capacity for zinc by Iranian clay, in their in-vitro tests.

In the case of hypokalemia, some in-vitro tests have been conducted by Gonzalezet a! .

{1982)56 showing a moderate potassium-binding capacity of clay especially at pH 6.

They suggested that clay eaters could develop hypokalemia depending upon the type of

clay ingested, the daily dietary potassium intake, and the renal function status.

Severance et a/ (1988)162 reported that the potassium level returned to normal when the

clay ingestion was discontinued and potassium supplement was given162• Gelfandet a/

(1975)53, analysing hyperkalemia in five patients with chronic renal failure, suggested

that since river-bed clay contains as much as 100 meq of potassium in I 00 g of clay

(much of which is exchangeable at acid pH), hyperkalemia appears to be the result of

the absorption of potassium released from clay after ingestion. As in the hypokalemia

case, the hyperkalemia ceased to be a problem when the patients stopped eating clay53•

One may ask, after reading the medical literature, if the clay (and more specifically the

clay as in termitaria) is the cause of certain mineral deficiencies or is it the cure? Most

probably it will depend on the physiological state of the individual, but the composition

of the termite mounds (or clay) eaten is of prime importance. Most of the research to

date on human geophagy has concentrated on clays or earths. The mineral content of

the clays studied, as indicated by the literature review is generally very low in

comparison to the high mineral content of termite mounds.

20

1.2 Nutritional Aspects of Selected Elements and Recommended Dietary Intakes

(RDis).

The traditional medicinal uses of termite mounds, such as during pregnancy and for

gastro-enteric disorders, would suggest, as previous authors have already mentioned

(chapter 1 . 1 .4), that elements such as Ca, Cu, Fe, Mg, K, Na and Zn may be of some

importance. In order to determine the nutritional contribution of termite mounds towards

the human nutritional needs, a summary of the selected elements nutritional

characteristics is a prerequisite.

The selected elements studied in this thesis have been classified in human nutrition as:

- electrolytes (sodium, potassium),

- major minerals (calcium and magnesium),

- trace elements (cobalt, copper, iron, manganese and zinc),

- substances with no known essential nutrient function in man

(aluminium).

A summary of the selected element (calcium, cobalt, copper, iron, potassium,

magnesium, manganese, sodium and zinc) body contents, physiological functions,

Recommended Daily Intakes (RDis), sources and deficiency symptoms is given m

Table 1.2. In this study, mineral and element are used interchangeably.

The RDis have been defmed as "the levels of intake of essential nutrients considered,

in the judgement of the National Health and Medical Research Council, on the basis of

available scientific knowledge to be adequate to meet the known rmtritional needs of

practically all healthy people. "124 The human element requirement is difficult to

establish as it varies between individuals and changes according to age, environment and

physical condition. This has led to differences between various national and

international recommendations. However, these differences are, however narrowing as

more knowledge accumulates169• In Table 1.2, the Australian RDis have been chosen

21

whenever available; the American values were selected for cobalt, copper and

manganese.

The RDis should not be misinterpreted as a daily minimum or as a requirement for any

specific individual. The RDis exceed the actual nutrient requirements of practically all

healthy persons as the estimates of requirements for each age/sex category have been

increased by a generous factor to take into consideration the variations in absorption and

metabolism124• As discussed by Southgate et a! (1989Y69, the mineral nutritiona! value

of food and diets is not necessarily equal to their composition as determined by chemical

analyses. The intestinal absorption and subsequent metabolism of all the elements needs

to be considered. Only a proportion of the total ingested element is capable of being

used. The amount of element absorbed depends not only on the chemical form of the

mineral in the food but on the other ingredients in that food and of the rest of the diet

and also on physiological factors169• Certain food in the diet may enhance absorption

of a particular element and decrease another, for example: ascorbic acid and protein

from meat decrease copper absorption and increase iron absorption. The problems of

minerals are multiplied by their own interactions. For example, an excess of zinc, iron

or calcium decrease the absorption of copper (an increase of 5 to 20 mg of zinc results

in an increase of the copper need of 75%)19,

Iron deficiency is the most common nutrient deficiency disorder in the world125 and

pregnant women subgroup is one of the most at risk categories. The incidence of iron

deficiency anaemia among pregnant women varies from I 0% in adequately nourished

groups to 50% in poorly nourished groups with multiple, closely spaced pregnancy183.

The iron deficiency results from one or a combination of the following: inadequate diet,

impaired absorption, blood loss or repeated pregnancies20• The iron is absorbed in the

duodenum and upper jejunum through a complex but poorly understood process183• The

factors which determine the proportion of iron absorbed from food are complex. They

include the iron status of an individual, the iron content and the composition of a meal.

Only a small proportion of dietary iron is absorbed; normal subjects commonly absorb

5-10% of the iron of mixed diets, and iron-deficient individuals 15-20% or more of this

iron, but considerable divergence from these values can occurm. The iron absorption

22

is also markedly increased during pregnancy, being about 30% in the second trimester

and 40% in the third trimester183• The composition of a meal also determines what

proportion of its content can be absorbedm. The dietary factors enhancing the iron

absorption are : ascorbic acid, citric acid, meat, fish and alcohol while the inhibiting

factors are polyphenols (such as tannins), phosphates, bran, phytate (in cereals and

legumes) and cooked egg yolk169• All meat sources promote the absorption of iron from

other foods. Even relatively small quantities of meat and fish (50-75g) may markedly

improve the hie-availability of iron124• Concomitant doses of 200 mg of ascorbic acid

with iron salts may increase absorption by 25 to 50%119•

Adult males and post-menopausal women measured iron losses are about I mg/day

(Table 1.2). Additional iron losses associated with menstruation vary, and for 90% of

women it averaged at 1 .35 mg/day124• During the second and third trimesters of

pregnancy, 5-7 mg/day extra iron is necessary to provide for the large increase in the

blood volume of the mother and the foetal growth. In Australia, the composition of the

diet (particularly its content of vitamin C and meat protein) suggest an iron absorption

rate of 15-20%, and therefor� the minimum dietary iron intake necessary to meet the

physiological requirements of adult males and post-menopausal women is 7 mg. For

menstruating women, 12-16 mg/day are necessary.

All elements are potentially toxic in large dose. The RDis, even though known to be

excessive for at least 80% of the population, are also known to be safe for 100% of the

healthy population. Data which would permit the delineation of toxic levels of dietary

elements are meagre. The toxicity level of an element depends on the extent to which

other elements which affect their absorption and retention are present. This is

particularly true for copper. In animals, a particular level of copper intake can lead

either to copper deficiency or toxicity, depending on the relative intakes of molybdenum

and sulfur, or of zinc and iron184• Toxicity iron ingestion is the fourth most frequent

cause of poisoning of children in United States. The average toxic dose being 200 to

250 mg!Kg; an acute toxic dose may be as low as 150 mg!Kgm.

23

Aluminium has been selected not for its potential nutritive aspect but for its possible

toxicity. Aluminium has been considered to be a benign metal by many authors like

Davidson et a! (1973)39• They claimed that it is too insoluble in its natural form to be

absorb by the body and this very property gave them a use in human medicine. For

example, aluminium silicate (kaolin) is widely used as an absorbent in the treatment of

diarrhoea. In 1970, Berlyne et a/, cited in Kundu (1990)" reported that high levels of

aluminium in tap water used for renal dialysis equipment could be linked to a form of

dementia in patients who were undergoing treatment. Since then, aluminium as been

assodated with a variety of metabolic and neural dysfunctions. The extent of absorption

of aluminium from clay, however small, could be of importance. As it is believed that

a relatively small amount of aluminium enters through the mucosal lining of the mouth

and the gastro-intestinal wall. The great majority is excreted unabsorbed92. Brown

(1983), as reported by Kundu (!990)" has shown that while increased amounts of

aluminium associated with reduced pH is certainly toxic, small amount of calcium may

prevent the toxicity.

TABLE 1.2 Nutritional aspects ofselected elements: adult body content, physiological functions, deficiency symptoms, daily losses, o/o absorption from food, recommended dietary intakes (R.Dis) and sources.

Element

Cobalt

Copper

Calcium

lwn

Magnesium

Manganese

Potassium

Sodium

Zinc

Body content and primary concentration

1.1 mg Liver

80-120 mg Liver, brain, heart. kidneys 20.7 to 24.8 g per Kg of fat free body tissue1 99% of Ca deposited in bones

4-5 g1u 70% in haemoglobin, 25% in storage form (ferritin, hemosiderin) Liver, spleen, bone marrow

20-28 g165 55% in bone, 27% in musculature 12-20 mg Kidneys, pancreas, liver

J I .S to 131 g1SO Principal intracellular cation 83-97 giSO major cation in extracellular fluid

1-2.3 g in almost aU tissue, muscle 63%, bone 20o/o, blood 2%1n

Data from 601 otherwise specihed.

Physiological functions Deficiency Symptoms

Integral part of vitamin 812 Pernicious anaemia

Haemoglobin synthesis, bone Anaemia, growth retardation. mineralisation, enzyme function secondary iron deficiency

Bones and teeth, blood Osteoporosis, poor coagulation, storage and release developme'nt of teeth and of hormones, activation of bones, delayed coagulation enzymes systems Transport and utilisation of Anaemia, disturbance of bone oxygen, component of energy marrow function transfer oxidase

Cofactor in enzyme reactions Neuromuscular disturbances, behaviour disturbances, cardiac disturbance

Bones structure, reproduction, Impaired growth, skeletal activator of enzyme function abnormalities, ataxia,

convulsions, vomiting Osmotic pressure Lethargy and tetany

Osmose regulation, water balance Dehydration, nausea, anorexia, fatigue, muscular cramps

Metalloenzyme function, protein Poor growth and sexual metabolism, lipid metabolism development, impaired wound

healing

Daily losses in mglday flo absorption]

[30"/o]

12�100-150 [20%]

124Men, pmw®: I Menstruating women: I.35 pregnancy: 5·,. [15-20%]

124[50%]

101[40%]

1241200

124140 + sweat (2· 8g/L)1SO

124 Adult: 2; Pregnant: +I; Lactation:+ 1.3 [30%]

RD!s for adults in mglday

m Supplied as vitamin 812, approximately 0.003

2-3

124 Adult: 800 Pregnant: + 300 Lactation: + 400

124Men, pmw®: 7 Menstruating women: 12-16 Pregnant: + 10-20

124Men: 320, women: 270 Pregnant: + 30 Lactation: + 70 Adult: 2.5·5

119Adult: ID-22

mAdult1950-5460

124Adult: 920-2300

124Adult: 1 2 Pregnant: + 4 Lactation: + 6

@: pmw post�menopausal women #: only for second and third tnmesters of pregnancy

Sources

Liver, meal Varies on soil content on which food is grown Nuts, shellfish, dried legumes, liver Milk. cheese, nuts, green leafY vegetables, bones

Liver, meat, oysters, nu"

Grains, fiuits, vegetables, nuts

Green leafY vegetables, nuts, grain, tea Fruits and vegetables

Sodium Chloride, meat, fish, milk eggs

Meat, liver, eggs, seafood

t

25

1.3 Some Aspect of Traditional Aboriginal Health Concept and Background

1.3.1 Traditional Aboriginal Health Concept

The concept of health and illness in the traditional Aboriginal world is quite different

in philosophy and practice to western medicine. The maintenance of health is tied to

spiritual, religious and social welfare123 " ••• rather than through adequate nutrition,

exercise and the maintenance of a hygienic environment1195• Serious illness and death

may be attributed either to sorcerers or to the effect, direct or mediated, of the breach

of a religious law or social norm. The most commonly postulated cause of illness is

sorcery153• Treatment with medicine is often secondary to the spiritual healing processes.

"However, in the case of minor ailments, from which the patient would be expected to

recover in any case, such as colds, gastric troubles, wounds or skin diseases, the

treatment would probably be with medicines only"114• Usually women are the herbal

medicine authorities in the community82•174, although every adult possess a

comprehensive knowledge of bush medicines123• The two methods of healing (spiritual

and medicinal) may be used in conjunction 174•

1.3.2 Aboriginal Health Background (Mineral Nutrition)

Franklin and White (199It9 reported that it is now generally accepted that the average

pre·colonial Aborigines were generally in good nutritional health. They were vigorous

people with few but robust children (a quarter of their children had died by the end of

their fifth year). Their life span was 40 years, with injury (including warfare and

murder) being the most frequent cause of death before disease. In general they ate well

and "their environment provided ... food comprising both protein and vegetable foods

with adequate vitamins and minerals. "49 "There was no evidence of rickets or other

nutritionally·related disease in traditional Aboriginal groups"95• "Early data from

recently nomadic Aboriginals generally indicated high haemoglobin concentrations for

both men and women"95• But "the rapid shift to a sedentary life on cattle stations, . . . ,

26

had a dramatic effect on the nutritional and health status of Aboriginal people"9s. Their

transitional diet indicated that sufficient iron was still usually provided but many diets

were inadequate in calcium, vitamin A and vitamin C95• T o·day, malnutrition appears

to be a persi_stent problem for many Aborigines112• "Aborigines and Torres Strait

Islanders comprise the least healthy identifiable sub-population in Australia . . . death

rates are up to four times higher, and life expectancy is up to 21 years less"182•

"Intestinal infections and infestations remain a major cause of Aboriginal ill-health and

of hospitalisation. Dia"hoea/ diseases were responsible for more than 10% of the

deaths of Aboriginal children aged less than 5 years in the NT (1979-1983}"182• There

has been an increase in anaemia, in more recent studies, mainly due to iron deficiency95,

" ... nutrients such as zinc, vitamin C, vitamin D, iron, and vitamin A, may be major

contributing factors to poor health in Aboriginal children"95• But, in other areas, Lee

(1991)95 reported a surprisingly large ranges of both red blood cell and serum thiamine

concentrations, including some high values on remote aboriginal communities.

1.4 Termitaria Biological Background

1.4.1 Taxonomy and General Biology of Termites.

27

Termites are polymorphic eusocial insects comprising the order Isoptera. They belong

to the superclass Hexapoda and the infraclass Pterygota (Figure 1.2). They are closely

related to the order Blattodea (cockroaches)89'91• This small group of primitive insects

contains about 2300 species world-wide with some 350 species in Australia192 and about

60 species in the Top End of Australia113• The lsoptera order has five families present

in Australia (Figure 1.3) with the Termitidae (higher termites) being the largest and most

recent52• Termites are one of the predominant groups of tropical invertebrates86,85• They

are found mainly in the tropical and subtropical regions of the world54•97•9, approximately

between 45"N and 45CS96, In Australia, termites are absent from certain types of soil207

and there is a very small number of species in rainforest areas 51•

Termites are popularly known as White ants'. This name was given to them by the

English in the West Indies and later used by early naturalists (eg: Sir Joseph Banks in his journal (1768-1771) cited in Watson and Gay (1983)"1• This taxonomic

misconception is still used today and often people confuse termites {lsoptera) with ants

(Hymenoptera).

Their physical and social characteristics are among the factOrs that influence the features

of the colony. Termites are soft-bodied insects with cryptic habits. They are

hemimetabolous (have no pupa stage) and they live in family groups (colonies) with

polymorphic castes. Three major castes are recognised: reproductives, workers and

soldiers (shown in Figure 1 .4) The ratio of the castes may vary with time; it is kept in

control through pheromones, hormones and selective cannibalism176•

28

Phylum ARTHROPODA

I Sub-Phylum ANTENNATA

I Super-Class HEXAPODA

I Class INSECTA

I Sub-Class D!CONDILIA

I CERCOFILATA

I Infra-Class PTERYGOTA

I NEOPTERA

I DICTYOPTERA

Order I BLATTODEA I I ISOPTERA

FIGURE 1.2 Classification of Isoptera (redraw from Kristensen, 199 191).

29

I ORDER ISOPTERA I I

I I Primitive Recent

(no worker caste, (worker caste) pseudergates)

I I RlllNOTERMITIDAE TERMITIDAE Coptotermes Amitermes

Nasutitermes Tumu/itermes

IMASTOTERMITIDAEI I TERMOPSIDAE I KALOTERMITIDAE I -

FIGURE 1 .3 Tennite Families (redraw from Hadlington, 198764)

The primary reproductives , or kings and queens, are derived from the alates151• In all

but one species, Mastotermes darwiniensis, the fore and hind wings are similar, therefore

their name !so( same) ptera(wings). After the nuptial flight their wings are shed and only

some small scales remain (Figure 1.5). The alates are fully matured insects with

compound eyes, hard pigmented cuticle and gonads. Their function in the colony is

reproduction. In many species the queen' s abdomen becomes distended due, mainly, to

the enlargement of the ovaries. This phenomenon is known as physogastry (Figure 1 .6).

The queen can lay up to 2000-3000 eggs per day192 and can measure up to 12 cm106•

Both king and queen live for many years, often over twenty. In some species, the

colony may survive the death of their progenitors by producing supplementary

reproductives or neotenics192•

30

' I

\\\ ,\,

- .'\' :;··· . ·- _:· I ' . \ ;''' . ··/ l,i' .;,.;,_� � y

\\ - I ' !'

. .

B

2·5.m m

c

'�

FIGURE 1.4

Castes of Coptotermes acinaciformis, A, winged reproductive or alate; B, worker; C, soldier. (from Watson & Gay, 1991 192 )

'l,'-A� I mm

FIGURE 1.5

De-alate reproductive (dropped wings) revealing severed wing butts. (from Hadlington, 198764 )

31

The workers class is the most abundant. They perform all the duties except defence and

reproduction. Their development is arrested at an early stage. They are sterile and

wingless and their compound eyes and ocelli are absent or greatly reduced 127• Their

cuticle is thin and highly susceptible to desiccation.

The soldiers class is the most distinctive with their head greatly developed. There are

two physically and different types of soldiers: the mandibulates and nasutes (Figure 1 .7).

Each species has one or the other type. The mandibulates (such as Amitermes vitiosus

and Coptotermes acinaciformis) have large characteristic jaws at the front of their head,

while the nasutes (such as Nasutitermes triodiae and Tumulitermes pastinator) have their

head ending in a long rostrum or nasus which opens in the front. Through the nasus

they can eject a sticky secretion onto their enemies112• The soldier's role is solely

defence. The males and females are sterile and usually apterous and blind. '

) )

. ' . .

' ' (\

•

A

' �::

l .. i ' · · ! 1 m m

FIGURE 1.6 Queen termite with a distended abdomen (physogastric queen) (from Hadlington 198764)

FIGURE !.7 A, mandibulate soldier of Heterotermes ferox, Rhinotermitidae; B, nasute soldier of Nasutitermes exitiosus, Termitidae. (from Watson, 1991192)

32

1.4.1.1 Feeding Habits

T errnites feed almost entirely on materials rich in lignin and carbohydrate, especially

cellulose118•190, Their diet ranges from living vegetation (trees, grasses, roots), to sound

dead wood, to decaying or even rotten plant materials206•118•29•190• Some species feed on

dried animal dung48, hurnus104 or soil rich in organic matter 97•206•85•111.29• Fungi are an

important part of the diet of many termites206'97'158'116• Cannibalism is also frequent in

termites not only as a means of sanitation206 and control of the population but also as a

means of conserving and recycling nitrogen113 and other minerals126• Termites can also

fix nitrogen by gut symbionts31'161.2°3•

The digestion of cellulose is facilitated by enzymes (including cellulases) secreted by

symbiotic Protozoa in the hindgut of the lower termites or bacteria in Tennitidae93• The

habit of oral and proctodaea! feeding of one individual by another helps to spread the

symbiotic protozoa and bacteria through the cornmunity129• Cellulases are also secreted

in the midgut of termites" (Figure 1.8).

.�..,Mouth

FIGURE 1.8 Digestive tube system in worker termites. (from Park, 1989139)

The workers are the only caste to gather food. Most of them consume their food in situ,

but many species (harvesters) carry it back to the nesf2•97"192 and stack the vegetative

material in certain chambers designed as attics127 or may build mounds which serve

largely as food stores97•206• lbis enables termites to operate throughout the extensive dry

season4• The dependent castes (soldiers, young larvae, nymphs and reproductives) feed

solely from the workers.

33

1.4.2 Nests

T ennites build a great variety of nests. They may be mere networks of simple galleries

in wood or soil, or elaborate constructions below and/or above ground level85• The nests

are usually characteristic of the species and in some cases of the genus64• Harris (1961 )68

remarkS that the species in residence is not always the original builder . Some species

(inquilinists) prefer to settle in the mound of another speciesm·127•23•193. According to

Noirot (1970)127, as many as ten different termite species can be found in a few cubic

decimetres of a Cubitermes built mound.

The function of the nest is to 'homeostatically' regulate several factors (e.g., humidity,

temperature, entrance of intruders, invasion by fungi)68•176•23• It may also, as mentioned

previously, serve as a food store97; Wood 1978; Watson 1991). From the nest, termites

may construct a system of galleries and covered runways which are known to extend -

from 50-75 m in Coptotermes and Nasutitermes up to over 100 m inMastotermer2• The

network can cover as much as a hectare�' and up to two hectares for some African

termites26• Some nests may extend several meters below the surface. Yakushev, quoted

in Lee and Wood (197la)97, reported termite galleries in West Africa going to the water

table at a depth of 70 m.

About 20% of Australian termites build nests in the form of mounds192 and Braithwaite

(1990?8 indicates that a third of the Top End termite species build termitaria (termite

mounds). The mounds vary greatly in size, shape and colour according to the species

of their builders, the environment and feeding habits62,611?7·'68• They can be a spectacular

feature of many savanna landscapes85• Their colour varies from grey to reddish-brown

depending on the material used for their construction57. Termitaria include by far the

largest structures built by insects. They range from hardened pavements to the 1cathedral1

structures of Nasutitermes triodiae which can extend up to 7 m high129•192• The diameter

can vary from a few centimetres to about 60 m96• Intraspecific variation in mound

structure can occur in some species72,tSt,97•1 56•147•93• Variation may range from 'no-mound

formation' to as many as five different types of mounds. In the same area, within a few

kilometres, different types of mound construction of the same species can be seen156•

34

It is therefore not always possible to identify a termite species from the size and shape

of the mound alone147•

The structure of the mound varies greatly from species to species. Noirot127 describes

two general types of mounds: the homogeneous mounds (all the chambers are alike and

there are no differences between the peripheral and internal regions) and the

heterogeneous mounds where different regions (outer soil capping, wall and nursery)

have different functions and may be composed of different materials (Figure 1.9).

The distribution of termite mounds is mainly affected by the environment167•102• It is

highly correlated with vegetation57 and there seems to be an inverse relation between the

termitaria and the soil nutrients97•145•136•57• Their density is very variable; it ranges from

less than 5/ha5 for mounds supporting very populous colonies (more than 1 million

termites per mound) to more than I 000/ha for small mounds208•

" !

; I ��-.•. - . 1.( , -, . ,( ; . ( . ' ' ·' <',;, � .

: ; --, .._ ·�,-�:: -� . . .

A

FIGURE 1.9 Mound nest:

Hdrd and thick

·�

Mo!(' open but hard inner layer

' Subterranean tunnels-In search ol lood

8

A: mound nest of Coptotermes /acteus B: mound nest of Coptotermes /acteus showing -internal structure (from Hadlington, 198r)

35

1.4.2.1 Mound Construction

The mound is not a static structure, it is in constant evolution and transformation26•97•

Basically there are two ways of building nests: excavation and construction. Excavation

is perhaps the most primitive and simplest action176• The construction techniques are

more sophisticated and may give rise to complicated nests in the higher termites.

Noirot (1970Y27 recognises two different methods of construction: addition and

modification. In the first instance new structures are added without any modification

of the existing ones (e.g., Cubitermes jungifaber) while in the latter, the workers modify

and re-organise the pre-existing structures by reworking the inside outwards (e.g.,

Macrotermes bellicosus, Nasutitermes exitiosus).

construction methods are combined127•

Very often, excavation and

Construction usually occurs at nighf2.23 and is usually seasonal. Most of the species

construct at the beginning205 or during the rainy season60•22 and a few others when both

the rainfall and temperature are low10•

1.4.2.2 Age of the Termitaria

There is a great variation in mound longevity, although little data have been collected

on this subject9•13•102• Mound longevity is generally based on the life-span of the colony,

since an unoccupied mound will erode more rapidly102 but in some cases, larger termite

mounds may be used by a succeeding colony of the same species after the first one has

become moribund147• Since some species are polycarpic with a single colony comprising

up to eight mounds80, the growth of the mound is not always related to the size and

environment of the colony. As a general rule, the growth rate of the individual mound

declines as the mound increases in height10•

The Australian termite mounds are ephemeral structures, most of them lasting for only

a few years, although the very large mounds of Nasutitermes triodiae may last 100

years or more96• Small colonies may reach their maximum size at the age of 4 and a

I I ,

!

36

half years and die at the age of 6-7 years, while larger colonies may die after 15-16

years 97• Holt et a! (1980f8 give a life-span of someAmitermes vitiosus mounds of 25-

50 years. The oldest termite mound known is described by Watson (1967)195 in Rhodesia

and may be 700 years old.

1.4.2.3 Termite Nest Material and Fabrics

Mounds are constructed from a mixture of materials such as soil, excreta, saliva and

plant debris206•68•97 combined into different fabrics according to the proportion of each

material. The choice of material depends partly on the termite feeding habits168 the

availability of material in their habitat68,97 and the part of mound being built (nursery,

wall, outer cap). Lee and Wood (197la)" identified two major types of fabric based on

the materials from which they were composed:

(A) Fabrics dominated by orally transported soil particles :

a) cemented solely with saliva as in the African Macrotermitinae mounds67•

b) re-packed with excreta, in addition to saliva. This type of fabric can be

found largely in the outer wall of mounds of a few species such as

Tumulitermes pastinato?'.

(B) Fabrics dominated by excreta:

a) consisting of soil (i.e. from soil-feeding species) mixed with orally

transported soil particles

,b) consisting of organic matter derived from the ingestion of plant material.

Sleeman and Brewer (1 972)168 observed that grass-feeding species (e.g. Nasutitermes

triodiae, Tumulitermes pastinator) appeared to build mounds composed mainly of soil

materials with little organic matter, while wood-feeders (e.g.Coptotermes acinaciformis)

tended to build mounds containing mainly organic material.

In general the same species seem to construct particular parts of a mound with similar

types of materials97• However, the proportions of the different elements may change

considerably with location68.97•98•168•

I

37

1.4.2.3.1 Soil

As discussed previously (1.2), soil particles are often the major constituents of the nest·

systems. Exceptions are found in the nest·systems of species which have no contact

with the soil, or in certain regions of the nests of other species.

A) Mound Material Origin and Clay Mineralogy

The origin of the soil particles used for the construction of termitaria has been discussed

by many authors and the clay mineralogy in mounds and soils has been used to identify ·

the soil material from which the mound originated98• Kaolin and illite represent the

major constituents of clay in the mounds and soils studied by Lee and Wood (197Ib)98,

They range from 60-100% of the total clay content. The kaolin alone varied from 10

to > 80% with a meao of 54% (Table 1.3). Boyer (1971)26 also showed high

percentages of kaolin in African termite mounds (70-85%). Generally, the kaolin ratio

is greater in the subsoil (B horizon) than in the topsoil (A horizon)98.

Sometimes building materials are brought from great depth; Boyer (1971 )26 found that

Macrotermes subhyalinus brought building materials from depths of 12-15 m. Lee and

Wood (1971b)98 have shown that the soil used by termites for their mound construction

generally comes from sub-soil. Many authors working in various parts of Africa

204,197,194,137,109,104,6s,61.26,134,144,71, India84,166,152,s6, British Guianas' and Australia<Js,2os have come

to the same conclusions. There are few exceptions, with a few studies reporting the use

of topsoil in mound construction98·102·209·104·136•9·144• Work reported by Lee and Wood

. (1971 b )98 show that in many places the topsoil contains too little clay to meet the needs

of mound construction, while the subsoil seems to satisfy requirements98• In their study,

based on 12 species taken from 1 7 localities in the Northern Territory, they showed that

half of the mounds studied came entirely from subsoil and the minority (20%) from

topsoil. The remainder derived from a combination of the two. They also showed that

38

a particular species does not always take soil from the same horizon. In four different

sites, Coptotermes acinaciformis used the B horizon twice, the A horizon once and in

the fourth site, a mixture of the two horizons. Nasutitermes triodiae used the same B

horizon in all the sites examined. They also noted that at a same site, mounds of several

species sampled showed that the materials came from different horizons. For example,

at one site, Tumulitermes pastinator andAmitermes laurensis used soil material from the

A horizon while Nasutitermes triodiae and Drepanotermes rubriceps used B horizon

material.

B) Particle Size.

The particle size of soils can be classified into the following categories: clay <.002 mm,

silt 0.002-.02 mm, fine sand 0.02-0.2 mm, coarse sand 0.2-2.0 mm and gravel >2.00

mm2• The size of the workers seems to dictate the upper limit of the grain size that can

be transported by mandibles or ingested97•109• Comparatively large species of termites

like Drepanotermes rubriceps and Nasutitermes triodiae incorporate a small fraction ( 1-

3%) of fine gravel in their construction72•98,

The clay fraction is an important element of the mound; it is not only a binding material,

but also helps to hold moisture in the mound. Three major reviews97•109•102 have been

published on the particle size of tennite mounds and their associated soils with emphasis

on the effects of termites on soil properties. The majority of studies show an increase

in the clay content of the mounds in comparison with unmodified soils. Although a few

authors have reported a similar clay content compared to the surrounding soil1ss,t49•6 or

subsoil166•71 or even lower clay content than the subsoW8•144•98• Differences in clay

content may occur within the mound. Some researchers have reported higher clay

content in the queen's chamber and the nursery71•7.26, while others found higher values

in the surface layer and outer wall 5°. Miedema and Van Vuure (1977)109 have questioned

the validity of some particle size analyses, as they detected some erroneous results of

particle size analyses. These are probably due to substances incorporated by the termites

in their mounds which could interfere with the soil fractionation. For the same reason,

39

Lee and Wood (1971b)98 have omitted physical analyses and detennination of clay

minerals of samples with high carton content (wood feeding termites).

Certain African termites may include up to 70% clay in certain structures of their

mounds24 and most of the mound-building species studied in Australia incorporate more

than 20% of clay in their structures (Table 1.3). Although the clay content of the

mounds can be up to 20% more than in any of the soil horizons, many species appear

to have no precise requirements of particle size98• They seem to be able to construct

mounds from a wide range of available materials. For example, two mounds of

Amitermes meridionalis located 100 m apart had different physical compositions: clay:

13 and 17%; fine sand: 42 and 27-29% respectively. With the exception of the inner

part of the mounds, the particle size composition of the nests seems to be more related

to the composition of the soil than the species of termite97. This contrasts with some

African species (e.g., Apicotermes occultus Silvestri) whose tu1dergrotu1d nests always

consist of 8 1·83% of fine sand mixed with organic matter, irrespective of soil type1n.

1.4.2.3.2 Saliva

The use of saliva as a binding agent for the construction of the nest and associated

structures has been noted by many authors6s,127,97,98,26,55,m,w7,5o,J32,149,

m.n.25·147·129·96. Its use may be more important for species which do not use their excreta

as a cementing agent97·98. Little is known about its composition. The saliva sustains

larvae, ftu1ctional reproductives and soldiers of some species. Lafage and Nutting

(1978)93 observed that the salivary gland foam cells secrete lipids and perhaps

mucopolysaccharides, and the vacuole cells, a substance with glycoprotein characteristics.

Grasse and Gharagozlou (1963)59 considered the saliva to be very rich in proteins and

Cmelick (1971, 1972)34·35 suggested the presence of some free fatty acids and easily

emulsifiable phospholipids.

40

1.4.2.3.3 Excreta

Termites use of excreta as a binding agent for soil particles has been reported by many

scientists96.Js.m,to7•93•36.84,9S,209,9•102• Lafage and Nutting (1978)93 noted that probably all

termites use their excreta for construction, modifying and lining their nests and

associated structures. Lee and Wood (1971a)97 observed that excreta can also be used

to fill disused galleries or as a structural component. The quantity and composition of

the excrement varies with the feeding habits and the structure considered. The soil�

feeding termites produce excreta which contains a large proportion of soil and appears

very much like soil. Wood-feeding species, in contrast to grass-feeding species, produce

a greater quantity of excreta. In these species, it may be the major constituent of their

nest or certain parts of it. This material is generally known as carton. It is a hard, dark

brown and highly adaptable construction material. It is made up principally of organic

excreta with some mineral particles and some fragments of undigested plant tissue911•93•

It contains up to 16.5% of polyphenolic material which resembles the alkali-soluble,

acid-insoluble humic materials of soils98• Lignin is a major constituent of carton,

although it also includes som� cellulose and sometimes substances that inhibit microbial

and fungal growth". Samples of cartons have been found to contain 87-95% of organic

matter (dry weight)".

1.4.2.3.4 Plant Remains

Some species such as Coptofermes, Nasutitermes and Mastotermes, have been observed

to incorporate some undigested wood into their mounds. This appears to be

exceptional'27•97• Likewise, the inclusion of grass into predominantly earthen mounds

seems to be incidental. Harvested plant material is generally stored within the galleries

and chambers of the mound.

41

1.4.2.4 Chemical Analyses

Much work around the world has been reported on the chemical analyses of termite

mounds and their surrounding soils. The aims of many of these studies was to assess

the influence of termites on biological, ecological or pedological factors. Therefore the

analyses concentrated on plant nutrients (organic carbon, total nitrogen, pH, phosphorus,

potassium, calcium, cation exchange capacity and exchangeable cations [calcium,

magnesium, potassium and sodium]). Very few studies have included other e�ements

(aluminium's·36•1•136; iron14•27•1•136; manganese36.136; zinc197'36). In Australia, only one study

undertaken by Okello-Oloya et a/ (1985)136 reported concentrations for aluminium and

manganese. The chemical analyses of termite mounds has been reviewed by Lee and

Wood (1971a)97, Boyer (1971)" and (1973)27, Bache1ier (1977)9, Wood and Sands

(1978)"" and more recently Lobry de Bruyn and Conacher (1991)'"'. Unfortunately the

species26,27, the age of the termitaria27 and the exact location of the sample (mound or

soil) is not always stated; the number of mounds investigated by researchers is often

small, sometimes only one, and an apparently unaffected soil may have been affected

by termite nests in the past102• According to Lee (1983)96 "much of the area of northern

Australia, and to a lesser extent of southern Australia, probably owes its present soil

mantle to termite-transported material". Work on chemical analyses of Australian

termite mounds have been conducted mostly in Northern Australia. The most

comprehensive study was done by Lee and Wood (1971b)" in 1971. Another major

research program was undertaken by CSIRO in North Eastern Australia to investigate

the role of termites in the ecosystem136,13s,no,t73•172.171•

Most of the studies, in Australia and throughout the world, showed an increase in the

chemical composition (organic carbon, nitrogen and exchangeable bases: mainly calcium

and magnesium) of termite mounds compared to the adjacent soiJI4.98·n,t02• However, a

few authors such as Nye (1955)134, Lee and Wood (197la&b)'�', Kang (1978)17, Sheikh

and Kayani (1 982)'63, and Spain and Mcivor (1988)171 observed that not all termite

mound materials are richer in plant-nutrients than the surface soils distant from the

mounds. Nye (1955)134 found that the chemical composition of the mounds seems to

42

vary according to the species of termite, the age of the mound, the part of it sampled

and the type of soil. Boyer (1956)25 in Central Africa reported a gradual increase of the

total bases content from the soil, to pediment, to wall and finally to the nursery. Those

differences are also seen clearly in the work by Lee and Wood (1971 a&b)"''· In

contrast, Pomeroy (1976)'44 in Uganda found no significant differences between new and

old parts of the mound, or between inner and outer parts of the mound wall. He

attributes this to the redistribution throughout the mound by diffusion during the wet

seasons. Coventry et a! (1988)38 in Australia, working onAmitermes vitiosus, came to

the same conclusion when they reported no significant difference between sampling

positions.

The cause of the mineral accumulation in termite mounds can be attributed to three

major factors:

I) the differential selection of soil particles and the use of a richer sub-soil by

termites201

II) the incorporation of vegetation, saliva and excreta (in certain parts of the

mounds)

III) pedological changes198•

Other factors which may contribute to mineral accumulation are:

a) the higher level of microflora activity in the mound. Cellulose decomposers142

and denitrifier micro-organisms (Pomeroy 1983)145 release nutrients into the

mound by their activity

b) the higher evaporation from the soil surface of the mound201•144•145•71•27 which is

increased by the air flow through the mound26.201•202• However, Fyfe and Gay

(1938)50 reported the presence of a relatively impermeable surface layer which

impeded evaporation in Nasutitermes exitiosus mounds

c) Watson (1976)199 reported the 'umbrella effect' of termite mounds in shedding

rainfall and retarding leaching in low and medium rainfall zones. He also

showed that water was retained through the wet season in termite mounds but

was leached from adjacent soils196

43

d) Another cause was proposed by Boyer (1973)27 who suggested that the

mechanical action of termites (grinding and remoulding soil materials with

vegetation) may free certain elements.

Although it is not always possible to compare the chemical analyses data :

• different procedures give different results,

- the element analysed and the species studied vary from one investigation tO another,

- and the exact location of the sample in the mound is not always known:

a few tables of data for chemical analyses of Australian termites have been compiled for

comparison purposes (Tables 1.3 - 1 . 1 0).

Table 1.3 Australian termite mound and adjacent soil physical properties ... ... The letters: *A to *D indicate the references.

Location Termite species Type of material Gravel C. Sand F Sand Silt Clay Kaolin Illite Others & & % % % ••

Reference Position Texture (excluding gravel), in % % NT Howard Spring Coptotennes acinaciformis M. outer casing 0 2 1 4 1 5 3 1 >80 . 10-20

*A A (0·25) <I 3 1 46 6 1 6 >80 . 5-10 8(60-75) <I 34 43 3 2 1 >80 . 1·5

N.T. S.Port Darwin Amitenne.s meridiana/is upper mound galleries 0 27 42 1 1 1 3 >80 . 5-10

*A M. center 0 34 42 10 1 3 >80 . 5-10 soil beneath mound 7 40 3 9 9 1 1 >80 . 5-10

Amitermes meridiana/is upper mound galleries 0 2 1 27 1 6 27 >80 . 5-10 M. center 0 29 29 1 1 27 >80 . 5-10 soil beneath mound <I 29 30 12 26 >80 . 5·10 A l l (0·10) 5 47 43 4 5 >80 . 5·10 Al2(10·25) 1 8 47 42 3 6 >80 . 5-10 B (25-45) 70 47 4 1 3 7 >80 - 5-10

N.T. Daly River Nasutitermes triodiae M. outer galleries 0 1 8 4 1 1 2 25 65-80 20-30 1·5 *A nursery 0 9 3 7 1 9 29 65-80 20-30 <I

basal region of mound <I 1 3 46 1 2 23 65-80 20-30 1-5 AI (0-6) <I 23 54 14 9 40-50 30-40 10-20 A2(10-20) I 22 52 12 1 3 30-40 50-65 1-5 B (40-50) 35 20 45 1 4 1 8 65-80 20-30 1-5

N.T. Pine Creek Coptotermes acinaciformis M. outer casing 0 1 1 39 20 25 50-65 30-40

'A Tumulitennes hastilis Mound 0 1 1 3 3 26 26 30-40 50-65 A I (0-10) 9 1 8 56 24 5 40-50 50-65 B (20-40) 49 IO 37 24 28 40-50 40-50 1-5

N.T. Larrimah Tumulitennes hastilis Mound galleries 0 3 8 22 7 29 >80 - 10-20 *A galleries under mound 2 34 25 3 37 >80 - 10-20

Amitennes vitiosus Mound galleries 0 33 30 4 29 >80 - 10-20 Mound base I 4 1 30 3 26 >80 - 10-20

AI (0-6) [ 46 32 9 8 >80 - 10-20 B (25-30) 2 36 1 9 4 44 >80 - 10-20

N.T. Tennant Creek Drepanotermes rubriceps Mound external wall [ 14 59 3 20 50-65 30-40 2-10 'A Mound internal [ [5 60 5 20 40-50 20-30 l-5

A (0- 10) [ 1 9 65 4 9 40-50 20-30 15-30 B (20-30) <[ 1 7 63 2 16 50-65 30-40 l l-25

QLD Mareeba I Amitermes laurensis Mound 0 12 41 25 17 10-20 65-80 10-20 'A Tumulitermes pastinator Mound outer galleries 0 4 45 27 1 9 10-20 65-80 10-20

Nasutitermes triodiae Mound outer galleries <[ 12 38 20 25 10-20 50-65 20-40 Drepanotermes rubriceps Mound outer galleries 3 1 0 43 22 26 10-20 50-65 25-50

A (0-8) <[ 12 56 23 7 10-20 50-65 5-10 B (16-24) <[ 7 43 20 29 20-30 65-80 15-30

QLD Mareeba 2 Amitermes /aurensis Mound outer galleries 0 [4 44 4 34 65-80 20-30 <[ 'A core of mound 0 1 9 49 5 24 65-80 20-30 <[

Nasutitermes triodiae Mound outer gaJieries 0 18 47 6 25 65-80 20-30 l-5 basal region of mound [ 1 9 47 5 23 65-80 20-30 <[ A (0-20) 1 8 26 65 4 4 40-50 40-50 <[ B (45-60) 4 1 27 40 7 24 50-65 30-40 <[

QLD Mareeba 3 Coptotermes acinaciformis M. outer casing 0 6 1 7 2 53 65-80 - 20-30 'A G. within wood 0 5 9 45 69 65-80 - 20-30

Schedorhinotermes int.act. G. over log 0 6 10 18 64 65-80 - 20-30 Nasutitermes triodiae Mound outer galleries <[ 30 [6 15 36 50-65 10-20 20-30

A (0-10) [ 8 18 24 53 65-80 - 20-30 B (30-50) 0 5 10 15 7 1 65-80 - 20-30

QLD Atherton Nasutitermes magnus Mound outer galleries [ 15 33 22 26 40-50 40-50 5-10 'A A (0-15) 7 15 39 30 15 20-30 50-65 5-10

B (25-35) 4 l3 35 26 25 40-50 40-50 6-15 QLD Townsville Amitermes laurensis Mound outer galleries 0 7 4 [ 27 19 50-65 10-20 2()..40

'A core of mound 0 6 42 29 20 50-65 10-20 20-40 Coptotermes acinaciformis M. outer casing 0 l3 45 18 19 50-65 10-20 15-30 Nasutitermes longipennis Mound 2 1 7 46 18 1 6 50-65 10-20 20-40

cont ... .. ....

Table 1.3 (cont ..• ) Austalian termite mound and adjacent soil physical properties ... "'

Location Tennite species Type of material Gravel C. Sand F Sand Silt Clay Kaolin Illite Others & & % % % ..

Reference Position Texture (excluding gravel), in % % B (16-24) I I 7 28 23 40 65-80 5-10 15-30

QLD Warwick Nasutitermes magnus Mound outer galleries I 52 21 9 16 >80 . 10·20

'A AI (0·20) 4 54 24 I I 10 >80 . 10-20 A2 (40-60) I I 50 23 9 17 >80 . 10-20

N.S.W. Canberra Nasutitermes exitiosus outer soil cap 0 26 3 1 9 30 50-65 20-30 1 1-25

'A Coptotermes /actus outer soil cap 0 16 26 7 47 50-65 20-30 10·20 AIA2 (0-20) 29 35 47 10 9 40-50 40-50 5·10 A3 (25-45) 44 36 43 10 I I 40-50 40-50 5-10 B (55-70) 38 24 28 9 40 65-80 10-20 1·5

N.S.W. Tallangatta Coptotermes /actus outer soil cap 0 27 13 17 37 20-30 40-50

'A AI (0·12) 6 32 26 19 22 20-30 40-50 A2 (15030) 4 26 28 17 28 40-50 10-20 B (35-55) 3 24 28 15 3 1 20-30 20-30 1·5

N.S.W. Narrandera Drepanotermes rubriceps surface galleries <I 22 52 9 14 30-40 40-50 1 1 -25

'A deeper gal. 5-30cm <I 24 52 8 14 30-40 40-50 1 1 -25 soil under ga11.30-60 I 25 54 6 14 30-40 40-50 1 1-25 soil below gall.60-70 4 23 54 7 15 30-40 40-50 1 1-25 • (0·30) <I 25 52 8 14 30-40 30-40 1 1·25 • (30-60) I 26 54 6 14 30-40 30-40 12·30 • (60-90) <I 19 54 7 23 30-40 40-50 12·30

S.A. Chowilla Drepanotermes rubriceps surface galleries . 4 1 38 9 I I

'A deeper ga1.2.5-7 .Scm - 38 34 10 16 g.bottom nest7.5-15cm . 37 34 9 17 soil- (0-2.5) . 51 35 6 7

- (2.5·7.5) . 45 39 7 7

• (7.5-15) . 44 40 8 7

• (15-20) . 39 34 7 1 8

TABLE 1.3 (conti.) Austalian termite mound and adjacent soil physical properties Location Termite species Type of material Grace I C. Sand F Sand Silt Clay

& & % Reference Position Texture (excluding gravel}, in %

'F QLD Amitermes Jaurensis Upper mound 21

Charles Towers Middle mound 24

Lower mound 20

Mound pediment 17

In term. soil (0·1 O)cm 8

*F N.S.W. Nasutitermes exitiosus Surface layer 17.7 43.0 17.0 22.4

Canberra Outer wall 16.8 41.1 16.7 25.4

Inner wall 26.7 37.8 19.3 16.3

Nursery 60.6 23.5 10.9 5.0

Soil 22.6 50.2 18.2 9.0

Nasutitermes exitiosus Surface layer 28.1 44.8 16.0 1 1 .2 Outer wall 33.9 41.0 15.2 10.0

Inner wall 36.8 37.4 18.1 7.7

Nursery 58.4 25.7 1 1 .3 4.6

Soil 33.2 45.3 15.3 6.2

Nasutitermes exitiosus Surface layer 21.1 33.7 16.0 292

Outer wall 16.7 33.6 16.1 33.6 Inner wall 23.8 34.1 21.4 20.6

Nursery 54.3 19.8 12.3 13.6

Soil 30.5 42.5 18.3 8.6

Nasutitermes exitiosus Surface layer 18.3 29.6 14.4 37.7

Outer wall 17.6 30.5 14.9 37.1

Inner wall 27.6 28.6 19.4 24.5

Nursery 58.1 25.0 4.0 12.9

Soil 39.6 38.0 16.4 6.1 n; Montmonllmute, Venmcuhte, Chlonde, regularly mterstratihed mateHai, ra!ldomly strat1hed matenal, quartz, haemahte, goeih11e and felspar. •A: Lee & Wood 1971b91 [Method Hutton (1955)] •G: Spain etal 1982'""' [Wt% of fine earth] H: Ho1t eta/ 198071 [Wt% of fine earth] •G: Spain et al 1982''" (WI o/e of fine earth] •F: Fyfe & Gay 1938"' (Method of Prescott & Piper 1928, after ignition]

Fyfe and Gay resuhs were re<:alculated to obtain the fractions (clay, silt, fine sand and coarse sand) in %

... 00 Kaolin % Illite % Others

.. %

TABLE 1.4 l\feans and standard deviations for a number of species of Australian termite mounds (and corresponding soils) chemical analyses

The !etten • A to •p indicate the reference

•• soil : fraction in the range of0-10 em depth

Reference • pll Org. C K,O K Ca p Mg Na Fe

g/lOOg mg/lOOg mgllOOg mg/lOOg mg/lOOg mg/lOOg mg/100g mg/lOOg

Tz£e of material mem SD mean SD mem SD mem SD mean SD mean SD mem SD mem SD mean SD

*A Mound: OM<IO% (n-39) 5.6 0.7 2.8 1.6 1699 1362 183 137 79 5 1 1 6 1 7

Mound: OM>lO% (n=26) 4.2 0.7 30.8 14.5 190 127 3 1 5 !54 24 10

Mound (n=65) 5.0 1.0 14.0 16.6 186 132 173 !56 19 15

•• Soil (n=18) 5.9 0.6 1.4 1.5 2290 1605 148 142 60 56 1 1 6

*B Mound 6.2## 0.5 2.1## 1.4 1045# 1 184 10.2# 6.5 15.9# 2.2 18.1# 7.2 241# 289 1372# 138 ( #o n� 6) (##o n�l8)

* * Soil (n=6) 5.9 0.6 0.8 0.2 1103 1455 8.2 4.5 1 1 .4 4.1 1 1 .6 3.2 254 257 1 132 143

*C+*D Mound (n=5) 5.8 0.2 1.9 1.0

** Soil (n=3) 6.0 0.2 0.6 0.1

*E Mound (n=2) 6.9 0.0 1.9 15.0 7.1

** Soil (n=2) 6.8 0.1 1.1 15.0 7.1

*F Mound (n=2) !53 19 135 44 135 30 5.3 0.7 1772 629

** Soil (n=2) 83 17 10.5 0.7 33 7.8 3.2 0.1 2561 2518

Mound mean (total) 5.3 1.0 10.8 15.1 1699 1362 256 412 !59 !54 18.5 14.1 47 55 182 267 1472 323 (n max = 92)

** Soil mean (total) 6.0 0.6 1.2 1.2 2290 1605 363 780 45 52 1 1 .5 5.5 1 7 1 0 191 246 1489 1165 (n max = 31)

•A: Lee and Wood (197lb/' n= number of data set from literature values

•s: OkeUo-Oloya et al. (1985)130 #: n= 6 •c: Holt and Coventry (1981)77 + Coventry i!l al. (1988)" ##: ll"' 18

•E: Birkill (1985)10 OM: Organic matter .... *F: Barr et al. (1988)14 "'

TABLE 1.4 (conti.) Means and standard deviations for a number of species of australian termite mounds (and corresponding soils) chemical analyses "'

0

The letters • A to *E indicate the reference ** soil : fraction in the range of 0-1 0 em depth

Reference * Mn A! Ext.P Exc.Ca Exc.K Exc.Mg Exc.Na S Tot. S Sol.

mg/IOOg mg/IOOg mg/IOOg mg/lOOg mg/lOOg mg/IOOg mgltOOg mgllOO mg/100 g g

Type of material m<m SD m<m SD rn<m SD mem SD mean SD mem SD m<m SD m<m m<m

*A Mound OM:<J0%(n-39) 75 5 1 23 !5 29 !5 4.3 4.7

Mound OM:> 10% (n=26)

Mound {n=65)

Soil {n=l8) 49 5 1 !6 !6 !2 9 !.7 0.9

*B Mound 26.4# 5.2 3613# 483 3.3## 3.7 !39## 63 2!.5## 9.3 24.!## I J .4 10.1# !4.2 ( #, n� 6) (##: n=18

Soil (n=6) 24.7 4.8 3093 937 0.6 0.4 16.6 8.5 19.0 4.7 13.9 72 2.9 2.0

*C+*D Mound (n=5) !.0 0.4 !32 9! 1 1.1 4.0 53 !3.8 0.5 0.!

Soil (n=3) 0.5 0.2 40 23 6.0 !.5 28.5 !!.0 0.4 0.4

*E Mound (n=2) 0.5 0.2 28! 2! 228 12.4 280 32 9.2 3.3 20.0 !.!4

Soil (n=2) 0.3 0.2 !70 44 !51 47 !86 !3 8.0 !.6

Mound mean (total) 26.4 5.2 3613 483 2.6 3.3 !04 7! 28 39 37 47 5.0 6.6 20.0 !.!4 (n max = 90)

Soil mean (total) 24.7 4.8 3093 937 0.5 0.3 50 55 25 38 26 45 2.3 2.! 10.0 0.98 (n max = 29)

*A: Lee & Wood (197lb)"' n= number of data set from literature values *B: Okello-Oloya et al (1985) #: n= 6 *C: Holt & Coventry (1981) + Coventry el al. (1988) 11#; � 18 •n: Birkm (1985) OM: Organic matter

TABLE 1.5 Termite mound and soil chemical data I (Lee and Wood, 1971 b)98• II samples containing high level of organic matter pretreated with peroxide

Location Termite species I Type of material

NT Berrimah

NT Howard Spring

NT S.Port Darwin

NT Daly River

NT Pine Creek

Soil

Mastolermes darwiniensis Carton in wooden build. #

Amitermes sp. Galleries und. bark #

Coptotermes acinaciformis M. outer casing

Carton from mound # Microcerotermes nervosus whole mound #

soil soil: A (0�25)

soil: 0(60-75)

Amitermes meridionalis

soil

Amilermes meridionalis

upper mound galleries

M. center

beneath mound

upper mound galleries

M. center

soil beneath mound

Microcerotermes nervosus Whole mound # soil A I I (0-10)

8 (25-45)

Nasulllermes triodiae M. outer galleries

nursery # basal region of mound

soil A I (0-6)

8 (40-50)


Carton from mound # Nursery from mound #

Tumu/itermes hastilis Mound

soil A I (0-10)

B (20-40)

pH

5.2

4.1

4.8

3.2

5.3

5.2

5.3

5.5

5.8

5.1

5.6

5.5

4.7

4.9

5.3

5.4

5.6

5.6

6.2

5.8

5.7

4.7

3.2

3.2

4.9

5.8

5.6

Org. C

g/100g

46.0

10.0

2.7

43.0

1 1.0

3.4

0.2

4.6

1.3

0.7

6.3

1.9

1.1

14.0

0.6

0.2

2.7

w:o

2.5

0.5

0.2

2.4

43.0

50.0

4.6

0.5

0.4

K,O

mg/IOOg

100

1 1 0

80

490

430

450

330

370

320

570

490

1730

1820

2020

2950

1870

3100

4340

3820

2800

K Ca P Exc.Ca Exc.K Exc.Mg Exc.Na

mg!IOOg mg/IOOg mg/IOOg mgfiOOg mgftoOg mgllOOg mgfiOOg

65

24

21

23

48

7

6

28

36

20

64

74

54

100

14

21

230

280

200

60

170

260

1 10

1 1 0

250

160

490

560

70

20

1 1 0

170

1 0

<10

40

10

<10

40

10

<10

140

<10

<10

70

220

70

10

10

20

170

120

80

10

<10

21

7

9

16

25

9

4

10

8

5

1 8

1 6

1 1

25

4

9

1 1

27

I I

4

8

12

1 5

1 6

1 4

7

18

20.0

10.0

2.0

24.0

8.0

0.8

24.0

10.0

0.8

0.6

0.2

4.0

150.3

60.1

4.0

2.0

14.0

52.1

2.0

6.0

3.5

1.6

0.0

4.3

12.5

1.6

13.7

22.7

2.0

0.4

0.4

39.1

62.6

54.7

5.9

2.3

10.9

17.2

7.8

3.5

41.3

2.4

3.6

9.7

3.6

0.2

14.6

4.9

0.2

0.2

0.1

38.9

83.9

34.0

3.6

9.7

25.5

37.7

9.7

17.0

2.5

1.6

0.2

5.5

1.8

0.9

9.4

2.1

0.7

0.7

0.2

0.9

2.1

1.8

0.9

0.9

0.9

1.1

1.1

2.3

cont .•.

"' �

TABLE 1.5 (conti.) Lee and Wood (l97lb) termite mound and soil chemical data

II samples containing high level of organic matter pretreated with peroxide

Location Termite species/ Type of materiaJ

Soil

pH Org. C K,O

total

K

HCI

gi!OOg mgltOOg mgllOOg

NT Larrimalt

NT Tennant Creek

QLD Mareeba I

QLD Mareeba 2

QLD Mareeba 3


Amitermes vitiosus

soil

Drepanotermes rubriceps

soil

Mound galleries

Mound galleries

Mound base

AI (0·6)

B (25-30)

Mound external wall

Mound internal

A (0-to)

B (20-30)

Tumulitermes coma/us Mudgut in dead tree # Amilermes laurensis Mound

Tumulitermes pastinator Mound outer galleries

Nursery from mound # Nasutilermes triodiae Mound outer galleries

Drepanotermes rubriceps Mound outer galleries

soil A (0-8)

B (16-24)

Amitermes laurensis Mound outer galleries

core of mound

Nasutitermes triodiae Mound outer galleries

basal region of mound

soil A (0-20)

B (45-60)


Carton from mound # G. within wood

Schedorhinotermes int.act G. over log

5.3

5.6

6.2

5.9

6.0

5.6

B 6.4

6.5

4.5

5.2

B 5.2

S.l

5.2

B 65

7.1

5.5

6.1

6.0

6.6

6.8

5.7

3.1

5.3

5.8

2.4

2.2

0.8

1.1

0.3

1.4

1.1

0.5

0.2

18.0

2.9

3.7

1 1.0,

4.4

1.6

0.8

0.4

2.1

1.3

2.0

4.0

0.5

0.3

2.7

44.0

2.8

7.4

370

350

350

490

370

1340

1310

1360

1430

4580

4340

4530

4840

5720

4790

1400

1420

1380

1440

3110

1510

220

180

260

74

64

53

61

64

170

170

130

160

260

180

250

270

310

260

100

380

120

98

140

150

38

160

100

54

94

97

c. HCI

mg!IOOg

60

80

50

60

30

130

90

70

40

170

60

90

260

70

30

<10

<10

100

70

50

160

30

20

90

300

40

210

� P Exc.Ca Exc.K Exc.Mg Exc.Na

HCI

mg!IOOg mgllOOg mgiiOOg mg/IOOg

13

14

13

14

13

I I

I I

10

8

1 8

I I

I I

22

10

8

8

8

16

14

12

12

8

20

10

20

9

I l l

64.1

62.1

38.1

36.1

26.1

114.2

92.2

60.1

42.1

50.1

84.2

68.1

24.0

2.0

2.0

1 14.2

64.1

88.2

142.3

30.1

38.1

164.3

104.2

240.5

15.2

12.5

9.4

6.6

6.6

19.6

23.5

14.5

11.7

5.9

11.3

7.4

5.1

2.0

2.7

16.4

10.2

23.9

46.9

5.5

11.3

39.1

34.4

32.1

19.5

26.8

14.6

12.2

12.2

21.9

10.9

10.9

9.7

34.0

64.4

64.4

55.9

8.5

79.0

32.8

12.2

31.6

47.4

4.9

18.2

42.6

36.5

34.0

mgiiOOg

0.7

1.1

2.5

1.4

0.7

1.1

1.8

0.7

0.5

4.6

25.3

17.9

9.0

1.8

10.6

3.7

1.6

1.4

3.4

0.7

0.9

3.4

2.8

3.9

QLD Atherton

QLD Townsville

QLD Warwick

NSW Canberra

NSW Tallangatta

Nas:utitermes triodiae

soil

Nasutitermes magrws

soil

Amitermes laurensis

Coptotermes acinaciformis

Nasulilermes longipennis

soil

Nasutitermes magnus

soil

Nasutitermes exitiosus

Coptotermes factus

soil

Coptotermes factus

Mound outer galleries A (0-10) B (30-50) Mound outer galleries Nursery of mound # A (0-15) n (25-35) Mound outer galleries core of mound M. outer casing Carton from mound # Mound A (0-8) B (16-24) Mound outer galleries Nursery from mound # AI (0-20) A2 (40-60) outer soil cap Carton from wall # Nursery from mound # outer soil cap Carton outer wall # Carton inner wall # Nursery from mound # AIA2 (0-20) Al (25-45) D (55-70) outer soil cap Carton from wall # Carton inner wall # Nursery from mound #

6.1 6.4 6.2 5.5 4.9 5.5 5.4 5.9 6.5 4.5 3.2 5.1 6.2 6.1 6.1 5.0 6.2 6.6 5.2 4.0 4.1 4.5 3.9 4.3 3.8 5.8 5.5 5.2 5.0 3.8 4.5 4.2

1.5 2.1 0.6 1.5

12.0 1.3 0.5 3.0 1.5 3.5

41.0 3.5 0.9

. 0.2 1.0

17.0 0.9 0.3 2.8

29.0 41.0

2.4, 29.0 41.0 42.0

1.6 0.3 0.3 4.8

44.0 43.0 50.0

710 270 220

2250

2950 2100 1640 1530 1470

1860 2720 1200

970

1000 960

1740

1540

2200 1710 2600

ISO 110 110 240 230 220 260

90 100 110 120

99 81

120 100 190

83 88

240 230 130 390 210 320 180 15 15

340 620 330 520 450

70 90 40 40

180 20 10

140 110 90

430 120

so 10 so

410 30 10 so

110 soo

40 310 520 400

90 30 10 70

390 480 400

39 14 9

19 27 18 17 10

6 5

18 8 6 3

16 48 18 16 13 29 39 1 4 2 4 47 31 I I

5 16 20 26 21 16

114.2 160.3

94.2 38.1

18.0 16.0

1 10.2 96.2 72.1

100.2 34.1 20.0 42.1

30.1 8.0

42.1

48.1

50.1 10.0 6.0

60.1

23.9 39.1 28.5 10.9

18.4 3.9

18.4 30.1 19.2

25.4 16.8 7.4 9.8

10.6 4.3

26.6

35.2

10.9 7.4

30.1 58.7

25.5 34.0 23.1

8.5

6.1 3.6

31.6 24.3 29.2

32.8 10.9 45.0 15.8

9.7 9.7

26.8

45.0

8.5 4.9

32.8 21.9

1.8 2.3 1.8 1.4

1.1 0.9 6.9 3.9 6.2

6.4 3.2

29.9 2.3

1.8 4.1 4.1

3.9

1.1 1.6 3.9 1.6

cont •..

l!l

TABLE 1.5 (conti.) Lee and Wood (1971b) termite mound and soil chemical data u. ..

II samples containing high level of organic matter pretreated with peroxide

Location Tennite species/ Type of material pH Org. C K,o K Ca p Exc.Ca Exc.K Exc.Mg Exc.Na

Soil total HCI HCI HCI

gllOOg mg!JOOg mg/IOOg mg!IOOg mg/IOOg mgi!OOg mgiiOOg mg!IOOg mg!IOOg

soil AI (0-12) 6.1 6.3 1940 490 230 2S 156.3 54.7 30.4 2.1

A2 (15030) S.8 !.6 1940 490 70 17 50.1 50.8 14.6 1.6

B (35-55) S.6 1.9 2020 S60 30 17 24.0 27.0 8.S 0.9

NSW Narrandera Drepanotermes rubriceps surface galleries 6.2 1.4 2240 JSO 170 21 156.3 50.8 21.9 3.0

soil • ( 0-30) 6.S 0.9 2470 320 130 17 122.2 39.1 17.0 2.8

- {30-60) S.6 0.2 2270 300 40 13 48.1 9.8 25.5 3.9

- (60-90) 7.1 0.2 2430 420 80 I I 84.2 39.1 65.7 25.3

SA Chowilla Drepanotermes rubriceps surface galleries 7.9 0.8 4SO 240 18 144.3 54.7 29.2 4.6

soil - ( 0-2.S) 6.9 0.4 320 80 13 74.1 43.0 20.7 2.S

- (2.5-7.5) 1.S 0.2 310 90 I I 62.1 32.1 20.7 3.7

• (7.5-1.5) 7.9 0.2 300 80 10 80.2 29.3 23.1 6.7

- ( 15-20) 8.0 0.2 430 90 12 74.1 26.6 43.8 48.3

- ( 25-30) 8.2 0.3 760 140 " 122.2 46.9 87.6 80.5

SA Gawler I Nasutitermes eJitiosus outer soil cap 4.3 4.0 1860 180 S2 ' 48.1 ,. 19.5 4h

Carton from wall N 3.9 15.0 116 JSO I I

Nursery from mound N 3.9 30.0 72 S20 19

soil AI (0-4) 4.9 !.S 2590 3 60 3 19.4 3.9 3.6 1.4

A2 (4-12) 4.S 0.4 2680 2 " 2 2.2 1.6 1.7 0.7

B (25-30) 4.3 o.s 1730 3 310 3 2.4 4.3 13.4 4.1

SA Gawler 2 Nasutitermes exitiosus outer soil cap 4.5 6.6 2930 530 100 16 1 12.2 16.4 45.0 7.8

Carton from wall N 4.0 26.0 238 440 24

Nursery from mound N 3.7 42.0 260 460 2S

soil AI (0-4) S.3 4.1 4380 380 170 13 146.3 12.5 23.1 S.3

A2 (6-12) 4.7 1.0 4660 400 15 13 26.1 4.7 8.S 2.1

B (25-30) s.o 0.8 3440 6SO 2S 18 42.1 12.9 38.9 S.l

TABLE 1.6 Coptotermes acinaciformis (Australian mound and soil chemical analyses)

Data from Lee and Wood (1971 b)"' II sample containing high level of organic matter, pretreated with peroxide

LOCATION TYPE OF MATERIAL pH Org. C K20 K Ca p Exc.Ca Exc.K Exc.Mg Exc.Na

POSITION total HCI HCI HCI

g!IOOg mg!IOOg mg!IOOg mg!IOOg mg!IOOg mg!IOOg mg!IOOg mg!IOOg mg!IOOg

NT Howard Spring M. outer casing 4.8 2.7 100 21 20 9 20.0 3.5 41.3 2.5

Carton from mound # 3.2 43.0 23 110 16

soil: A (0-25) 5.2 3.4 110 7 10 9 10.0 1.6 2.4 1.6

soil: 8(60-75) 5.3 0.2 80 6 10 4 2.0 0.0 3.6 0.2

NT Pine Creek M. outer casing 4.7 2.4 3100 260 20 12 14.0 10.9 25.5 0.9

Carton from mound # 32 43.0 1 10 170 15

Nursery from mound # 3.2 50.0 1 10 120 16

soil: AI (0-10) 5.8 0.5 3820 160 10 7 2.0 7.8 9.7 1.1

soil: B (20-40) 5.6 0.4 2800 490 10 18 6.0 3.5 17.0 2.3

QLD Mareeba 3 M. outer casing 5.7 2.7 220 100 90 10 164.3 39.1 42.6 3.4


soil: A (0-1 0) 6.4 2.1 270 110 90 14 160.3 39.1 34.0 2.3

soil: B (30-50) 6.2 0.6 220 110 40 9 94.2 28.5 23.1 1.8

QLD Townsville M. outer casing 4.5 3.5 1470 110 90 5 72.1 19.2 29.2 6.2


soil: A (0-8) 6.2 0.9 2720 81 50 6 34.1 16.8 10.9 3.2

soil: B (16-24) 6.1 0.2 1200 120 10 3 20.0 7.4 45.0 29.9

th "'

TABLE 1.7 Amitermes vitiosus (Australian mound and soil chemical analyses) II: mean mound value

Location Type of material pH Carbon p Ca K

g!IOOg mg/IOOg mgllOOg mg/IOOg

Mean S.D. Mean S.D. Mean S.D. Mean S.D. Mean S.D. ·Basalt Wall Upper mound 6.3 0.2 2.6 1.0

red earth Middle mound 6.4 0.2 2.1 1.2 15.3# 2.4# 5.2# 4.9# 200# 30#

•a Lower mound 6.6 0.4 I.S 0.6

Mound pediment 6.4 02 2.1 1 2 14.8 3.9 7.5 62 180 30

Soil (O·IO)cm 6.3 0.1 0.9 0.2 12.0 3.4 5.9 3.8 130 50

Basalt Wall Upper mound 6.4 0.2 1.8 1.0

yell. earth Middle mound 6.5 0.3 1.8 0.6 15.5# 3.8# 4.4# 1.6# 230# 40#

•a Lower mound 6.4 0.3 1.2 0.5

Mound pediment 6.4 0.1 1.4 0.5 15.1 3.3 6.1 3.8 200 20

Soil (0-IO)cm 6.3 0.2 0.9 0.2 14.2 1.5 7.7 4.2 170 20

Basalt Wall Upper mound 6.1 0.1 1.5 0.4

Grey earth Middle mound 6.4 0.2 1.3 0.3 12.5# 5.5# 6.8# 4.6# 290# 110#

•a Lower mound 6.0 0.3 1.3 0.3

Mound pediment 6.1 0.2 1.1 0.3 11.0 1.1 9.0 6.8 270 110

Soil (0-IO)crn 6.3 0.4 0.6 0.1 8.> 2.1 3.9 1.9 260 90

NT Larrimah Mound galleries 5.6 2.2 14 80 64

Lee & Wood Mound base 6.2 0.8 13 50 53

'A soil: A I (0-6) 5.9 1.1 14 60 61

soil: B (25-30) 6.0 0.3 13 30 64

Manbullo Mound 6.85 10.0

control Below mound 0-10 em 6.95 10.0

Birkill Below mound 10-30 em 6.95 10.0

'E Below mound 30-50 ern 6.90 10.0

Soil 0-10 em 6.87 10.0

Soil 10-30 em 6.85 10.0

Soil 30-50 em 6.75 10.0

Mg

mg/100g

Mean S.D.

10.2# 3.5#

1 1 .6 6.9 7.6 4.3

10.7# 4.4#

12.7 3.4

8.8 4.9

16.8# 7.2#

15.6 5.3

12.1 3.2

Na Fo mgiiOOg mgllOOg

Mean

7.0#

8.0 9.0

13.0#

39.0 13.0

23.0#

20.0 180

S.D. Moan

3.0# 1510#

4.0 1250

5.0 1210

2.0# 1440#

65.0 1290

6.0 1340

9.0# 1270#

14.0 l l 80 260 1190

S.D.

350#

330 550

110#

160

180

190#

90 200

"' a,

Manbullo

unfertilized 'E

Charters

Towers Holt

•o

Charters Towers

•o

ChartersTow.

•c

Mound

Below mound 0-10 em

Below mound 10-30 em Below mound 30-50 em

Soil 0-10 em

Soil 10-30 em

Soil 30-50 em

Mound

Below mound to 10 em Below mound to 40 em Soil 0-10 em

Soil 10-20 em

Soil 20-30 em

Soil 30-40 em Soil 40-50 em Soil 50-60 em

Soil 60-70 em

Soil 70-80 em

Mound Below mound to 10 em Below mound to 40 em Soil 0-10 em

Soil I 0-20 em

Soil 20-30 em Soil 30-40 em Soil 40-50 em

Soil 50-60 em

Soil 60-70 em

Soil 70-80 em

Mound n�to

below m. 0-40 em

Soil 0-10 em n=l8

6.90 1.91 20.0

6.62 1.17 20.0

6.47 0.64 20.0

6.52 0.32 20.0

6.72 1.09 20.0

6.60 0.61 20.0

6.80 0.39 10.0

5.9 1.13 - 1.24 - 0.63

5.8 0.65

5.8 0.34

5.8 0.29

5.1 0.28

5.1 0.21

5.8 0.17

5.8 0.13

5.8 0.13

5.6 1.28

- 0.78 - 0.56

5.9 0.65

5.1 0.34

5.6 0.29

5.6 0.28

5.5 0.21

5.5 0.17

5.6 0.13

5.1 0.13

5.9 2.23

6.2 0.58

6.2 0.55 *A: Lee & Wood ((9116)"'; •B: Okello-oloya et a/ (1985)""; °C: Holt & COventry(\982)"; •D: Coventry et a/1988"; *E: B1rktll 1985"'

"' ....

TABLE 1.7 cont.. Amitermes vitiosus (Australian mound and soil chemical analyses) #: mean mound value

Location

Basalt Wall red earth 'B

Basalt Wall yell. earth

'B

Basalt Wall Grey earth

'B

NT Larrimah 'A

Manbullo

control

'E

Type of material

Upper mound

Middle mound

Lower mound

Mound pediment

Soil (0-IO)cm

Upper mound Middle mound Lower mound

Mound pediment

Soil (0-IO)cm

Upper mound

Middle mound Lower mound

Mound pediment Soil (0-1 O)cm

Mound galleries Mound base

soil: A 1 (0-6)

soil: B (25-30)

Mound Below mound 0-10 em Below mound 10-30 em

Below mound 30-50 em Soil 0-10 em

Soil 10-30 em

Soil 30-50 em

Mn

mgl100g Mean S.D.

AI

mg!IOOg Mean S.D.

P acid-Extr.

mg!IOOg

Mean S.D.

Exch. Ca

mg!IOOg

Mean S.D.

Exch. Mg

mg/lOOg

Mean .D.

Exch. K

mg!IOOg

Mean S.D.

1.7 26.2# 7.7# 3510# 810# 2.0

2.0

0.7 156 34.1 25.5 1.5 145 25.9 22.6

1.2 156 44.5 16.8

7.9 27.0 10.9 5.4 25.8 8.6

4.3 22.3 10.9 24.5 26.5

6.0 3940 460 5.5 2110 1320

1.0 0.4 124 31.3 19.0 5.0 18.8 7.4 6.3 0.7 0.1 17 8.0 21.6 10.9 2l.l

1.9 I. 7 225 54.3 30.6 25.2# 7.1# 3960# 240# 1.9• l.3 211 67.9 29.4

24.6

33.3

6.6 4080 280 5.3 3610 180

1.9 0.8

0.6

l.3 209 54.5 18.2 0.2 150 53.5 21.3

0.2 21 5.0 15.7

18.9 4.3 3390 270 0.9 0.5 122 17.0 26.3

18.9# 4.3# 3390# 270# 1.0 0.5 141 44.1 27.4

0.9 0.4 163 55.7 23.0 19.5 2.9 3580 170 0.6 0.2 110 38.7 24.1 23.3 4.3 3390 350 0.3 0.2 8.2 2.6 22.3

0.70 0.50

0.50

0.50

0.50

0.50

0.50

62.1 38.1

36.1 26.1

295 204 166

141 138

148

123

26.8 14.6 12.2 12.2

302 175

165

165

177

205

168

6.7 23.1 7.3 30.5 4.1 28.5 6.1 21.5

2.2 23.1

3.5

9.4 5.1

4.3

2.7

2.8 20.3 4.3

6.3 26.2 6.3

5.8 32.8 8.2 4.5 22.3 5.9

4.3 25.0 6.3

12.5 9.4 6.6 6.6

237

237

178

127

1!7

99

80

Exch.Na

mg!IOOg

Mean SO

0.9 0.5 • • • •

0.7 0.2 0.7 0.5

1.8 0.5 • • • •

1.6 0.2

0.5 0.1

2.3 • •

1.8 5.3

1.1

2.5 1.4

0.7

11.5

9.2

9.2

9.2

9.2 9.1 9.2

1.8 •

•

1.1 6.9

S.Tot S.Sol

mg!IOOg mg/1 OOg

"' 00

0.00 Manbul\o Mound 0.38 266 258 219 6.9 20.0 1.14 unfertilized Below mound 0·1 0 em 0.17 202 200 182 6.9 10.0 0.92 'E Below mound I 0·30 em 0.10 148 170 141 6.9 10.0 0.69

Below mound 30·50 em 0.10 136 171 130 6.9 10.0 0.32 Soil 0-10 em 0.15 201 195 184 6.9 10.0 0.98 Soil 10-30 em 0.10 163 180 136 6.9 10.0 0.59 Soil 30-50 em 0.10 166 204 113 6.9 10.0 0.26

Charters Mound 0.90 56.91 70.13 8.99 Towers Below mound to I 0 em 1.30 67.53 49.11 12.12

Holt Below mound to 40 em 0.70 51.10 47.40 8.99 •o Soil 0·10 em 0.70 42.48 39.75 7.04 0.46

Soil 10-20 em 0.40 26.05 31.97 6.26 0.46 Soil 20·30 em 0.30 20.84 31.24 5.08 0.46 Soil 30·40 em 0.30 21.84 38.41 5.08 0.46 Soil 40-50 em 0.30 22.85 52.02 5.08 0.46 Soil 50-60 em 0.30 21.84 61.02 4.30 0.46 Soil 60-70 em 0.10 20.84 66.24 3.91 0.46 Soil 70-80 em 0.10 17.84 70.13 2.35 0.46

Charters Mound 0.60 42.48 62.96 5.08 Towers Below mound to I 0 em 0.90 58.12 53.48 10.17

•o Below mound to 40 em 0.60 40.88 49.84 7.04 Soil 0-10 em 0.40 16.�3 28.08 4.30 0.69 Soil 10-20 em 0.30 8.62 23.58 3.91 0.46 Soil 20-30 em 0.30 9.62 30.63 3.13 0.46 Soil 30-40 em 0.20 8.62 45.46 2.35 0.46 Soil 40-50 em 0.20 10.62 64.30 1.96 0.69 Soil 50-60 em 0.10 8.62 73.29 1.17 1.15 Soil 60-70 em 0.10 6.41 75.24 1.17 us Soil 70-80 em 0.10 4.41 79.74 1.17 1.84

Charters Tow. Mound n=IO 1.61 269.3 48.86 14.08 0.46 •c below m. 0-40 em n=40 0.62 92.18 19.45 6.65 0.69

Soil 0·10 em n=l8 0.48 61.72 17.75 6.65 0.23 "' 'D •A: Lee & Wood 1971"; *B: Okello-oloya et a/l98JD6; •t: Holt & Coventry (1982)11; *D: Coventry et all988"; *E: Birktll 19115'6

TABLE 1.8 Nasutitermes triodiae (Australian mound and soil chemical analyses) "' C>

Data from Lee and Wood (1971b)91 •BB: material entirely or predominantly derived from B horizon

Location Type of material pH Org. C K,O K Ca p Exc.Ca Exc.K Exc.Mg Exc.Na

total HCI HCI HCI

Possible source g/IOOg mg/IOOg mg/IOOg mg!IOOg mg/lOOg mg/IOOg mg!lOOg mg!lOOg mg!IOOg

NT Daly River M. outer galleries *BB 5.6 2.7 1730 230 70 1 1 4.0 39.1 38.9 0.9

nursery # 5.6 10.0 1820 280 220 27 150.3 62.6 83.9 2.1

basal region of mound 6.2 2.5 2020 200 70 1 1 60.1 54.7 34.0 1.8

AI (0-6) 5.8 0.5 2950 60 10 4 4.0 5.9 3.6 0.9

A2(10-20) 5.5 0.2 1970 84 <10 4 2.0 2.0 4.9 0.5

B (40-50) 5.7 0.2 1870 170 10 8 2.0 2.3 9.7 0.9

QLD Mareeba 1 M. outer galleries *BB 5.1 4.4 4530 310 70 10 68.1 7.4 64.4 17.9

A (0-8) 5.5 0.8 5720 100 10 8 2.0 2.0 8.5 1.8

B (16-24) 6.5 0.4 4790 380 10 8 2.0 2.7 79.0 10.6

QLD Mareeba 2 M. outer galleries *BB 6.1 2.0 1380 140 50 12 88.2 23.9 3 1.6 1.4

basal region of mound 6.0 4.0 1440 !50 160 12 142.3 46.9 47.4 3.4

A (0-20) 6.6 0.5 3110 38 30 8 30.1 5.5 4.9 0.7

B (45-60) 6.8 0.3 1510 160 20 20 38.1 11.3 18.2 0.9

QLD Mareeba 3 Mound outer galleries 6.1 1.5 710 !50 70 39 114.2 23.9 25.5 1.8

A (0-10) 6.4 2.1 270 110 90 14 160.3 39.1 34.0 2.3

B (30-50) 6.2 0.6 220 110 40 9 94.2 28.5 23.1 1.8 #: sample pretreated w11fl peroxtde (high organ1c carbon content)

TABLE 1.9

Data from

LOCATION

QLD Mareeba I

'A

Charters Tow

•o

Tumulitermes pastinator (Australian mound and soil chemical analyses)

•A Lee and Wood (1971b)"" •o Coventry et al (1988)'1

TYPE OF MATERIAL pH Carbon K20 K Ca p Exc.Ca Exc. K Exc.Mg Exc.Na

POSITION total HCI HCI

g/IOOg mg/IOOg mg/IOOg mg/IOOg mg/IOOg mg/IOOg mg/IOOg mg/IOOg mg/IOOg

Mound outer gaJieries 5.5 3.7 4340 250 90 I I 84.2 1 1.3 64.4 25.3

Nursery from mound # 5.2 1 1.0 270 260 22

soil: A (0-8) 5.5 0.8 5720 100 10 8 2.0 2.0 8.5 1.8

soil: B (16-24) 6.5 0.4 4790 380 <10 8 2.0 2.7 79.0 10.6

Mound n=9 5.6 3.6 0.94 141.48 12.90 50.32 0.69 below m. 0-20 em n=24 5.6 0.6 0.58 57.31 7.04 25.77 0.23

Soil 0-10 em n=IS 6.2 0.6 0.48 61.72 6.65 17.75 0.23

Mound Moan 5.6 3.7 4340 250 90 6.0 1 12.8 12.1 57.4 13.0 (SO) 0.1 0.1 7.1 40.5 1 . 1 10.0 17.4

Soil 0-10 em Mean 5.9 0.7 5720 100 10 4.2 31.9 4.3 13.1 1.0 (SO) 0.5 0.2 5.3 422 3.3 6.5 1 . 1

#: sample pretreared with peroxide (high organic carbon content)

"' �

"' "' TABLE 1.10 Tumulitermes hastilis ( Australian mound and soil chemical analyses) Data from •A Lee and Wood (197lb)91

LOCATION TYPE OF MATERIAL pH Carbon K20 K Ca p Exc.Ca Exc. K Exc.Mg Exc.Na

POSITION total HCI HCI

g/IOOg mg!IOOg mg!IOOg mg!IOOg mg!IOOg mg!IOOg mg/lOOg mg!IOOg mg!IOOg

NT Pine Creek Mound 4.9 4.6 4340 250 80 14 52.1 17.2 37.7 1.1 'A soil: A I (0-1 0) 5.8 0.5 3820 160 10 7 2.0 7.8 9.7 1.1

soil: B (20-40) 5.6 0.4 2800 490 <10 18 6.0 3.5 17.0 2.3

NT Larrimah Mound galleries 5.3 2.4 370 74 60 13 64.1 15.2 19.5 0.7

'A galleries under mound 6.4 0.6 370 62 40 14 40.1 9.0 12.2 1.6

soil: AI (0-6) 5.9 1.1 490 61 60 14 36.1 6.6 12.2 1.4

soil: B (25-30) 6.0 0.3 370 64 30 13 26.1 6.6 12.2 0.7

Mound M•an 5.1 3.5 2355 162 70 14 58.1 16.2 28.6 0.9

(SD) 0.3 1.6 2807 124 14 I 8.5 1.4 12.9 0.3

Soil 0-10 em Mean 5.8 0.8 1645 276 30 16 21.0 5.1 14.6 1.8 (SD) 0.2 0.5 1633 303 42 3 21.3 2.2 3.4 0.7

63

1.4.2.4.1 pH

The differences between soil and termite moilllds pH are generally small209•102•97 and seem

to have little slgnificance93• The increase of pH, mainly in Macrotermes spp. mounds,

seems to be linked to calcium carbonate accurnulation98•102, while their decreases in other

mounds may be related to the incorporation of organic-rich excreta209• The average

Australian molUlds and associated soils studied are acid in reaction: pH ranging from 4.2

• 6.9 in mounds with, lower values: 4.2 ± 0. 7 in the nursery or carton material and 5.9 -

6.8 in soils (Table 1 .4).

1.4.2.4.2 Organic Carbon

The organic content of most termite mounds is higher than the soil from which they

originated and occasionally lower than the surrounding soii97• It varies greatly not only

according to the species and the origin of the construction material (top- or sub-soil) but

also within the same mound26•97•98• In non-homogenous types of mounds, it generally

increases from the outer mound casing to the inner mound structures (carton or nursery).

For example, Coptotermes acinaciformis mounds have a mean of 2.8 ± 0.5 percent of

organic carbon in the mound outer casing and 44.2 ± 3.4 percent in the nursery and the

carton part (Table 6 (b)). In Australia, the average organic carbon values of mounds

composed largely of soils and their associated soils are respectively between 2 - 3

percent and around I percent (Table 1.4 (a)), while it reaches 50 percent in the nursery

area (Table 1.5 (a)). Okello-Oloya et al (1985)136 observed that the levels of carbon

vary within the mound and decrease from the upper to the lower levels. Organic carbon

also increases in the pediment compared with the soil.

64

1.4.2.4.3 Elements

A) Calcium

Nearly all the studies around the world showed an increase in the calcium content (total

or exchangeable) of termite mounds compared to the adjacent soils. In Africa and Asia,

nodules of calcium carbonate can be found in certain Macrotennitinae mmmds26.27•71•143•

In Australia no nodules have been found. Lee and Wood ( 197Ib)98 mention a usual

increase of calcium in tennitaria of two to five times compared to the soil from which

they originated. According to the Wood and Sands ( 1978)209 table an increase in

calcium (acid extractable) is not always followed by an increase of exchangeable

calcium, and vice versa. Some authors reported higher exchangeable calcium values

than acid extractable or total. There would appear to be some problem in their

analytical procedures. For example, in Okello-Oloya et a/ ( 1985)136, the total calcium

values range from 4.4 ± 1.6 to 21.8 ± 6.8 mg/1 OOg while the exchangeable calcium

values range from 122 ±17.7 to 227 ± 54.3 mg/IOOg (Table 1.7). The total calcium

values are abnormally low compared with the acid-extractable values reported by Lee

and Wood (197lb)" (Table 1.4). The procedure of extraction mentioned was a digestion

with hot hydrofluoric acid. While no specific details of the method are available, it is

likely that the fluorides would interfere with calcium extraction.

The increase of calcium content within mounds seems to follow the increase of organic

carbon content, although Lee and Wood (197la&b}97•98 see no precise correlation

between the two. Acid extractable calcium values in Australian mounds analysed by Lee

and Wood (197lb)" (Table 1.5) vary from <10 to 560 mg/100g with a mean of 173 ±

156 mg/lOOg in the mounds and 60 ± 56 mg/100g in the soils (0-10 em fraction or

closest to this profile ex.: 0-7.5 em) (Table 1.4). The exchangeable calcium values

follow the same kind of range (Table 1.4). The values vary greatly with the location,

species and the part of the mound analysed. For example, a Nasutitern1es triodiae

mound was found to have ISO mg/lOOg exchangeable calcium in the nursery and only

4 mg/1 OOg in the mound outer galleries (Table 1.5). Generally, Okello-Oloya et a/

( 1985)136 found in Amitermes spp. (Table 1.7) mounds that the exchangeable calcium is

65

higher in the upper and middle level part of the mound than in the lower and the

pediment section, but always higher than the surrounding soil.

B) Magnesium

Generally, the magnesium content of mounds is higher than that of the surrounding

surface soils7•14•27•26.25•21•132,14o,109•88•97•98, As for other elements, it varies with the species

responsible for the building204•163•98, or the location1s,!n,98• Most of the magnesium

reported in the literature is exchangeable magnesium. In Australia, total magnesium data

have only been reported by Okello-Oloya et a/ (1985)136 and Barr et a/. (1988)14•

Coventry et al (1988), reported an increase of exchangeable magnesium of 2.6 times in

Amitermes vitiosus mounds compared with the soil. Similar increases have been noted

for Drepanotermes and Tumuliterme�8• Lee and Wood (1�71 a&b)97�8 reported greater

increase of exchangeable magnesium for some species. For example, a Coptotermes

acinaciformis mound outer casing contained 41.3 mg/lOOg of exchangeable magnesium,

while its adjacent soil had only 2.4 mg/lOOg and a Nasutitermes triodiae mound had

83.9 mg!IOOg in its mound nursery compared with 3.6 mg/100g in the adjacent soil

(Table 1 .5). An increase of magnesium of 4-5 times has been recorded in two mounds

(traditionally used by Aboriginal people) compared to the surrounding soil14•

In Australia the average, total magnesium reported by Okello-Oloyaet al (1985)136 for

mounds of Amitermes vitiosus and Amitermes laurensis is 18 . 1 ± 7.2 mg/IOOg

(Table1 .4). This is comparatively low compared with the value they found for the

exchangeable magnesium 24.1 ± 1 1 .4 mg/100g (Table 1 .4). The reasons for this could

possibly be explained in the same way as already mentioned for calcium (sec 1.2.4.3 A).

The distribution of exchangeable magnesium in mounds and soils varies between sites

but seems to be consistently lower in the lower level of the mound136• The highest

values of magnesium were reported by Birkill ( 1985)21 in Katherine for A. vitiosus

mounds with a mean of 280 mg/1 OOg. She also reported high soil values, such as 186

mg!IOOg. The Australian average values of exchangeable magnesium in mounds are 37

66

± 47 mg/lOOg (Table 1 .4). These values are comparable to those reported for species

in other countries102•

C) Potassium

As with calcium, potassium figures in the termite mounds are generally higher than the

soil from which they originate14'98•136•38•98 but may be only slightly higher or even lower

than the topsoil figures209 and do not seem to be closely linked to organic matter (Table

1.4). As for calcium and magnesium they vary according to the species and the sample

position (see Table 1.5). There are greater differences between the total and the

exchangeable potassium than for calcium and magnesium. Okello-Oloyaet a! (1985)136

reported total potassium values ranging from 200 - 2720 mg/l OOg for mounds and 130 -

3520 mg/l OOg for adjacent soils, while the exchangeable potassium varied from 5.5 -

32.8 mg/100g to 13.3 - 25 mg/100g respectively. There are no special patterns of

exchangeable potassium associated with the different levels of mounds (upper, middle

and lower)136•

D) Sodium

As for the other elements already mentioned, sodium content seems to be higher in the

mound than in the surrounding soil, although Okello-Oloyael a/ (1985)'" showed that

at some sites the levels were much lower. Values vary greatly according to the

procedures used, the species and the location. Okello-Oloya et a/ (I 985)'" reported

total sodium values ranging from 7.0 ± 3.0 to 23.0 ± 9.0 mg/IOOg in mounds and 9.0

± 5.0 to 180 ± 260 mg/100g in the soil, compared with the exchangeable values of 0.9

± 0.5 to 2.3 ± 1.8 mg/100g in the mounds and 0.5 ± 0.1 to 5.3 ± 6.9 mg/100g in

adjacent soils (Table 1.7). Boyer (1956)" reported very high levels of sodium in the

inner part of the mound of Be!licositermes rex, probably due to the incorporation of

saliva during construction.

67

E) Phosphorus

In general mounds have a higher phosphorus content than adjacent soils (Table 1.4).

The average, dilute, acid-extractable phosphorus in termite mounds and soils studied in

Australian are respectively: 18.5 mg/IOOg and 1 1 .5 mg/IOOg. Coventry el a/ (1988)

reported values 2 to 3.7 times higher in the mounds. There are no consistent variations

between mou�d levels although the lower part of the mound seems to have a lower

content136• Lee and Wood {1971 a&b)97�8 reported that the distribution of phosphorus

in the mound is fairly uniform.

F) Iron and Aluminium

The iron and aluminium content of termite mounds has been poorly studied in Australia

and around the world. Stoops (1964)175 reported an increase of free iron inCubitermes

mounds. Boyer (1973)27 indicated values of 5.2 to 7.8 percent of Fl?:03 with an even

distribution through the mound of Bellicositermes spp. and 16 to 20 percent of Al203•

He also reported the presence of iron concretions27 and an accumulation of Fe2+ from

underground water which would eventually precipitate as ferric hydroxide. Agarval

(1978)1 reported a considerable increase in F�03 and Al203 in mounds compared with

the surrounding soil with not much difference between regions of the mounds, but he

did not differentiate between the iron and aluminium values. Cornaby and Knebs

(1975f6 reported lower aluminium values in the mounds than in the soil. The iron

values reported by Basalingappa et a/ (1978)15 in India are very similar to those of

Boyer (1973)27 in Africa, eg. 7.27- 7.91 %, with little variation within the mound. This

contrasts with Basalingappa et a/ (1978)u aluminium values which vary from 0.09% in

the mound (soil) and 14. 17% in the royal chamber. In Australia, Okello-Oloyael a/

( 1985)136 reported values for iron and aluminium respectively of 1 .4 and 3.6 % for

mounds and 1 . 1 and 3 . 1 % in the soils. This indicates a slight increase of those values

in the mound compared with the soil. The variations of iron between sites were also

very slight. This contrasts with the work of Barr et a/. (1988) in which high variations

of iron were reported between sites (Table 1 .4).

68

G) Manganese, Zinc, Cobalt and Copper

Very little has been reported on these elements. Comaby and Knebs (1975)36 in

Venezuela reported higher concentrations of manganese in the mound of Nasutitermes

sp. than in the nearby soil, while the zinc levels were lower. Okello-Oloyaet a/

(1985)136 reported manganese values of 26.4 mg/lOOg in the mound and 24.7 mg/lOOg

in the soil, while Watson ( 1970}'97 reported anomalous concentrations of zinc in termite

mounds compared with the surrounding soiL There have been no data reported on

cobalt.

1.4.2.5 Agricultural Uses of Termitaria

Termite mound use as a soil amendment has been discussed by many authors. The

results are often divergent. They depend on the properties of the crops, the soils and the

habits of different species of termitesn In deficient soils, termitaria can provide

nutrients7!). Sheppe (1968) observed that when the subsoil is richer than the topsoil, the

termitaria are used in preference to the adjacent soils by African natives to plant their

crops. In many parts of Africa and Asia, better crops such as vegetables133, sisal133•68•86,

sorghum148 maize147•103 cotton and tobacco68•86 have been obtained on termite mounds ' '

or fields where the termite mounds have been levelled. In Thailand, Pendleton (1941)143

reported that the farmers are using mounds for growing cotton, vegetables and tobacco

but that the productivity of the levelled termite mound is very irregular. In Australia,

Okello-Oloya and Spain (1986)135 have reported an increase of biomass of Digit aria

ciliris (an annual grass) and Stylosanthes hamata (a pasture legume) on termite mound

materials compared to surface soils from the same areas. The increase was correlated

to the phosphorus and nitrogen level of the mound and soil material used. Negative

results have also been reported by Nye (1955)134 and Kang (1978)", in Nigeria, where

growth of annual crops, such as maize was poorer in the soil mound or levelled mounds.

In Zaire, Meyer (1960i08 reported that when the material brought up by the termite from

the subsoil is particularly infertile, the mounds (mainly if they are abundant) may present

a serious obstacle to cultivation.

1.5 Aims of this Project

The aims of this project are to:

69

determine the elemental composition of termite mounds eaten by Aboriginal

people and more specifically, of two Aboriginal communities of the Northern

Territory, with the mineral analyses concentrating more particularly on iron,

calcium, potassium, sodium, magnesium, manganese, aluminium, copper, cobalt

and ziric.

determine the particle size fractionation of termite mound material

to assess the selected elemental variations:

I) between different age material (new and old)

2) within mounds according to the sample position: top, middle and bottom

3) between mounds of the same species (and different species) in a same

location

4) between mounds of the same species (and different species) in different

locations

determine the bio-availability (in vitro) of the selected elements studied (with

emphasis on iron bio-availability)

CHAPTER TWO

MATERIAL AND METHODS

•

2 MATERIAL AND METHODS

2.1 Collection of Termitaria

71

The method of sampling termitaria was designed as a result of consultations with

Aboriginal communities (section: 1 . 1.2.1 and 1 .1 .2.2).

2.1.2 Method of Collection

In Daly River, the surface samples (O-J em) were taken at random on the outside of the

tennite mound (on both new and old parts of the mound) using a stainless-steel knife.

The deeper samples (0-lOcm) were taken, randomly, at different height on the outside

of the motmd with a I 0 em core fixed on a cordless drill. Core samples were also taken

from the middle section of two mounds (Nasutitermes triodiae and Tumulitermes

pastinator). A minimum of 200-250g of sample was collected when possible.

In Elliott, the samples (approximately 10cm3) were cut using an axe, as used by the

Aboriginal people. All samples were stored in paper soil bags.

Ten centimetre cores of soil were collected, at each site, using a l Ocm auger. Triplicate

cores (50cm apart) were obtained for each sample site and bulked.

2.1.2 Site Locations

The samples were collected at 4 geographically different localities: Berrimah. Howard

Springs, Daly River and Elliott.

Geographical references concerning the different sites are given in Table 2.1.

;::! TABLE 2.1 Site locations

Site Termite species Location Longitude Latitude Grid reference Notes collected • (map)

Daly River (site I) Tp,l11 5070 Daly River 130" 43' 13" 44' 52LFK865803 5.2 Km on the right when coming from the Daly River mission

Daly River (site 2) Av 5070 Daly River 130" 42' 13' 45' 52LFK847785 3.4 Km on the left when coming from the Daly River mission

Daly River (site 3) Nt,Tp 5070 Daly River 130" 44' 13' 39' 52LFK883897 20.2 Km from the Police Station on the Daly River Road to Darwin

Daly River (site 4) Nt,Av,Ca 5070 Daly River 130' 49' 13" 3 1 ' 52LFL961 040 2.9 Km after Lichfield road when coming from the mission; 64.6 Km from Tipperary

Elliott (site 5) Av SE 53-5 N.waters 133' 30' 17" 35' 53KLA4055 8.4 Km on the left side of the road leading to Lake Woods and the Longreacb Waterhole

Howard Springs (site 6) Nt,Tp 5073-2 Darwin 130" 59' 12" 28' 52LGMI58207 0.5 Km after the Yarrawonga Zoo, on the left when coming from Darwin

Berrimah (site 7) Nl 5073-2 Danvin 130" 55' 12" 28' 52LGM093213 I .8Km before East Ann Settlement, on the left side of the road coming from Darwin

• Tp: Tunwlilermes pastinator Th: T111mditermes hastilis ca·: Coptotermes acinaciformis A v: Amitermes vitios11s Nt: Nasutitermes lriodiae

2.1.3 Sample Description and Summary

73

All soil and termite mound samples are numbered according to the following system:

Sample numbering system: Mound: AbcDe Soil: cDe

Ab: Termite species (Av: Amitermes vitiosus, Tp: Tumulitermes pastinator Nt:

Nasutitermes triodiae Th: Tumulitermes hastilis, Ca: Coptotermes

acinaciformis) c : Sample number

D : Indicates the geographic location (D: Daly River, E: Elliott, H: Howard Spring)

e : Site number in a specific location

An underlined sample is a collected on the outer surface of the mound (depth: 0-lcm).

An asterisk (*) at the end of a sample number indicates that the sample has been

collected on a newly built part of the mound by opposition to the "old'' material. Old

does not have a connotation of time (specific age)� it is a part of the mound material that

has been weathered. It is distinguished from the new built material by the different

texture and color.

Sample numbering example: Nt31D4* = Nasutitermes triodiae sample number 3 1 from

Daly River, site 4, collected on a new part on the outer surface of the mound (depth of

0-lcm).

2.1.3.1 Detailed Mound Study

One mound of each of the three major species used by the Aboriginal communities

(Nasutitermes triodiae, Tumulitermes pastinator andAmitermes vitiosus) was studied in

detail. The physical characteristics of the mounds are given in Table 2.2. A total of 20

samples were taken from Nasutitermes triodiae mound (Site 3), 17 from the

74

Tumulitermes pastinator mound (Site 3) and 9 from Amitermes vitiosus mound (site 5).

The different number of samples was related to the size of the mounds and the way the

Aboriginal people sample them. A detail of the 3 mound sampling is given in Table 2.3

and associated soil samples are given in Table 2.4.

TABLE 2.2 Physical characteristics of 3 termitaria selected for more detail sampling: Nasutitermes triodiae (Nt) from Daly River (site 3), Tumulitermes pastinator (Tp) from Daly River (site 3) and Amitermes vitiosus (Av) from Elliott (site 5).

Mound characteristics Termite species -

Nt Tp Av

Site number 3 3 5

Mound number I I I I

Basal circumference (em) 460 320 127

Middle height circum (em) 310 250 98

Top less lOcm circum (em) 72 130 53

Height (em) 310 70 64

Tota1 mound samples 20 17 9

Total soil samples 4 3 4

75

TABLE 2.3 Detail mound sample summary for 3 termitaria: Nasutitermes triodiae (Nt) from Daly River (site 3), Tumulitermes pastinator (Tp) from Daly River (site 3) and Amitermes vitiosus from Elliott (site 5).

Mound number/

0-lcm depth from outer casing

0-IOcm depth from outer casing

0-1 Ocm depth from middle section

site --------------------------'--_;_-

l / site 3

I / site 3

1 1 I site 5

Top Middle Bottom Top Middle Bottom Top Middle Bottom

NtOl "' Nt02

Nt04* Nt08* NtlO Ntl2 Nt05 Nt09 Ntll Ntl3

Nt03* Nt06*

Tp27',Tp28 Tp29,Tp30*

Tp31

Nt07

Tp32 Tp33 Tp34 Tp35

Tp36 Tp37

Ntl4 Ntl6 Ntl8 Ntl9 NtiS Nt17 Nt201

T38 Tp39

Tp40 Tp41 Tp42 Tp431

ns ns ns Av21 Av23 Av26 Av27 Av28 A29 Av22 Av24

Av25

•: newly built material 1: material from nursery ns: not sampled

TABLE 2.4 Soil samples collected at 0-!0cm depth in Daly River (site 3) and in Elliott (site 5).

Location

Daly River

Elliott

1-m away from any mounds

15D3, 16D3, 17D3, 18D3, 19D3

OlE, 02E

�1m away from any mounds

20D3, 21D3, 22D:l'

03E, 04E

76

2.1.3.2 Termitaria Sampling

In general three samples were taken per mound (Top, middle, bottom). If the mound

height was below !.2m, only 2 samples were taken (top and bottom). In the sample

summary tables (2.5, 2.6, 2.7, 2.8), the diameter + height gives an indication of the

relative volume of the mound. The circumference was taken at lm height for mounds

over l . m otherwise at half-height of the mound. All samples were 0-10 em depth,

unless indicated. The underlined samples were superficial samples (depth: 0-lcm). The

* indicates that the sample was taken on a newly built part.

2.1.3.2.1 Amitermes vitiosus Termitaria Sample Summary

A total of 1 0 Amitermes vitiosus mounds (5 small and 5 medium) were sampled in

Elliott, 14 mounds in Daly River: 1 0 mounds in Site 4 and 4 mounds in site 2, (Table

2.5).

2.1.3.2.2 Tumulitermes pastinator Termitaria Sample Summary

A total of 15 Tumulitermes pastinator mounds (5 large, 5 medium and 5 small) were

sampled in Daly River (site I) and 13 mounds (3 large, 5 medium and 5 small) in

Howard Springs (site 6) (Table 2.6).

2.1.3.2.3 Nasutitermes triodiae Te�mitaria Sample Summary

A total of 15 mounds (5 large, 5 medium and 5 small) were sampled in Daly River

(site4), 5 large mounds in Howard Springs (site 6}, and 5 large mounds in Berrimah (site

7) according to the availability of mounds in each location (Table 2. 7).

77

TABLE 2.5 Amitermes vitiosus termitaria sample summary

Mound Mound Height Circum Height + Sample location number/ class (em) (em) Diam (em)

Site Top Bottom

11 site 5 Sma11 23 32 33 AOl A02

21 site 5 Small 28 34 39 A03 A04

31 site 5 Small 28 50 44 AOS A06

41 site 5 Small 21 34 32 A07 AOS

51 site 5 Small 23 35 34 A09 A10

61 site 5 Medium 38 35 49 A l l A12

71 site 5 Medium 30 3 1 40 A13 Al4

81 site 5 Medium 35 45 49 A l 5 A16

91 site 5 Medium 45 70 67 A l 7 Al8

10/ site 5 Medium 30 55 48 Al9 A20

11 site 2 Small 75 87 103 A30 A32

21 site 2 Small 64 71 87 A33 A35

31 site 2 Small 57 89 85 A36 A38

41 site 2 Small 59 70 81 A39 A41

11 site 4 Medium 120 140 165 A42 A43

21 site 4 Medium 140 150 188 A44 A45

31 site 4 Medium 1 1 0 140 155 A46 A47

41 site 4 Medium 1 1 0 140 155 A48 A49

51 site 4 Medium 120 190 180 ASO ASl

61 site 4 Small 60 90 89 A 52 A53

11 site 4 Small 40 60 59 A 54 ASS

8/ site 4 Small 50 120 88 A 56 A57

91 site 4 Small 60 90 89 ASS A 59

10/ site 4 Small 60 1 1 0 95 A60 A61

78

TABLE 2.6 Tumulitermes pastinator termitaria sample summary

Mound Mound Height Circum Height + Sample location number/ class (ern) (ern) Diam (em)

Site Top Bottom

11 site 1 Large 80 340 188 TpOI T02

21 site 1 Large 90 330 195 Tp03 Tp04

31 site I Large 90 287 181 Tp05 Tp06

41 site 1 Large 100 454 245 Tp07 Tp08

51 site 1 Large 75 263 !59 Tp09 Tp!O

61 site I Medium 65 197 128 Tpl l Tp12.Tp13*

71 site 1 Medium 50 164 102 Tp14 Tp15

8/ site I Medium 54 261 137 Tp16 Tp17

91 site I Medium 60 200 124 Tp18 Tp19

10/ site I Medium 50 135 93 Tp20 Tp21

1 11 site I Small 17 60 36 Tp22 ns

12/ site 1 Small 21 58 39 Tp23 ns

13/ site 1 Small 18 61 37 Tp24 ns

14/ site 1 Small 20 75 44 Tp25 ns

15/ site I Small 10 40 23 Tp26 ns

II site 6 Medium 80 226 !52 Tp44 Tp45

21 site 6 Medium 75 203 140 Tp46 Tp47

3/ site 6 Medium 100 177 !56 Tp48 Tp49

41 site 6 Medium 75 377 195 Tp50 TpSI

51 site 6 Medium 60 206 126 Tp52 Tp53

61 site 6 Small 45 83 71 Tp54 Tp55

71 site 6 Small 28 39 40 Tp56 ns



I 01 site 6 Small 30 53 47 Tp59 ns

1 1/ site 6 Large I I 0 250 190 Tp60 Tp62

12/ site 6 Large 130 337 237 Tp63 Tp65

13/ site 6 Large 105 230 178 Tp66 Tp68

79

TABLE 2.7 Nasutitermes triodiae termitaria sample summary

Mound Mound Height Circum Height + Sample location number/ class (em) (em) Diam (em)

Site Top Middle Bottom

1 / site 4 Large 470 480 623 Nt21 * Nt22 Nt23

2 / site 4 Large 400 420 534 Nt24 Nt25 Nt26 Nt27,Nt30* Nt28,Nt31* Nt29,Nt32*

3 / site 4 Large 380 580 565 Nt33 Nt34 Nt35,Nt36* Nt37 Nt40* Nt38,Nt41* Nt39,Nt42*

4 / site 4 Large 370 480 523 Nt43 Nt44 Nt45 Nt46.Nt49"' Nt47Nt50* Nt48 Nt51*

5 / site 4 Large 380 670 593 Nt52 Nt53 Nt54 Nt55,Nt58* Nt56,Nt59* Nt57.Nt60*

6 / site 4 Medium 250 350 361 Nt61* Nt62 Nt63 Nt64 Nt67* Nt65,Nt68* Nt66,Nt69*

7/ site 4 Medium 190 240 266 Nt70* Nt71 Nt72

8 / site 4 Medium 240 280 329 Nt73 Nt7.4 Nt75

9 / site 4 Medium 260 460 406 Nt76 Nt77 Nt78

10 / site 4 Medium 250 500 409 Nt79 Nt80 Nt81

1 1 I site 4 Small 120 130 161 Nt82 ns Nt83

1 2 I site 4 Small 100 100 132 Nt84 ns Nt85*

13 I site 4 Small 100 180 157 Nt86 ns Nt87

14 / site 4 Small 1 10 100 142 Nt88 ns Nt89

1 5 / site 4 Sma11 120 120 158 Nt90 ns Nt91

I I site 6 Large 380 500 539 Nt92 Nt93* Nt94

2 1 site 6 Large 350 360 465 Nt95 Nt96 Nt97

3 1 site 6 Large 340 374 459 Nt98 Nt99 NtlOO

4 1 site 6 Large 370 474 521 NtlOJ Nt102 Nt103

51 site 6 Large 350 494 507 Nt104 Nt105 Ntl06

I I site 7 Large 440 580 625 ns Ntl07 ns

2 I site 7 Large 450 400 577 ns Ntl08 ns

3 I site 7 Large 470 420 604 ns Ntl09 ns

4 1 site 7 Large 550 470 700 ns NtllO ns

5 I site 7 Large 480 420 614 ns Nt1 1 1 ns

80

2.1.3.2.4 Tumulitermes hasli/is Sample Summary

One other species mounds was sampled at site 1 : Tumulitermes hasti/is. T. hastilis

mounds were widely distributed but not reported to be eaten by the aboriginal

communities of Daly River. Samples collected are shown in Table 2.8.

TABLE 2.8 Tumulitermes hastilis Sample Summary

Mound Height Circum Height + Sample location number/ (em) (em) Diam (em) Middle

site

I/ site I 90 86 78 ThO!

2/ site 1 45 52 47 Th02

31 site 1 50 44 82 Th03 4/ site 1 75 71 61 Th04

51 site I 87 64 52 Th05

2.1.3.3 Soil Sampling

A minimum of 3-4 soil samples were taken at each major site: Elliott (site 5), Howard

Springs (site 6), Berrimah (site 7) and Daly River (site I, site 2, site 3 and site 4). Each

sample was taken at a minimum distance of 1-m from any termite mound on the site and

at a depth of 0-lOcm. Another soil sample was also collected near site site 2 (50m on

the right to the Daly River mission) as it is favoured by Mercia' family during the

annual wet season flood, when the mounds are not available.

2.2 Sample Preparation

All the tennitaria and soil samples were initially crushed in plastic bags by gentle

rubbing with a piece of wood (the tennitaria often being in solid lumps). This allowed

the removal of the grass and tennites by sieving and shaking. The samples were then

81

dried, in soil paper bags, at 60° in an oven. The coarse sample was crushed using a soil

crusher (Conservation Commission Berrimah Soil laboratory) and sieved through a 2mm

sieve. A I OOg sub-sample of material was removed for physical analyses. The

remaining sample was pulverised using a ring grinder (<75 microns). A sub-sample of

the homogenous pulverised sample was taken for chemical analyses.

2.3 Particle-Size Analysis

The aim of the particle-size analysis was to subdivide the soil minerals into different

categories according to the particle diameter

clay <0.002 mm (<2 �m)

silt 0.002-0.02 rom (2-20 �m)

ftne sand 0.02-0.2 rom (20-200 �m)

coarse sand 0.2-2.0 mm (200-2000 �m)

The particle-size analysis method used was based on the Pipette and sieve method

described by Coventry and Fett (1979)37 in which sodium tripolyphosphate was replaced

by Calgon (hexametaphosphate) as dispersing agent. It was conducted at the NT Conservation Commission Berrimah soil laboratory. A chemical pretreatment of termite

mound samples rich in organic matter (>0.5%) with hydrogen peroxide was conducted

according to the method of Mcintyre and Loveday (1974)10s.

2.4 Acid Extraction of Termitaria and Soils

A number of digestion methods, using combinations of the acids HN03, HCl04 and

H2S04 were investigated. The aim was to select a method that would provide

reproducible results with high reproducible recovery rates. It was not necessary to

obtain a total extraction for this study. The elements analysed were aluminium, calcium,

cobalt, copper, iron, magnesium, manganese, potassium, sodium and zinc. The

82

determination of the element concentrations was performed using a Varian SpectrAA 40

atomic absorption spectrophotometer (AAS) (see 2.4.3).

2.4.1 Extraction Trials

The trials were conducted in order to select the most appropriate:

- acid combination (eg: nitric/perchloric (9:1), nitric/sulfuric (1 :4),

nitric/perchloric (1 :4)} for the extraction

- sample size (lg, 5g, lOg, 25g),

- sample fraction (2 mm, pulverised),

- type of digestion flask (150 mL beaker on hot plate, 150 or 200 mm test tubes

in a block digester).

The results of the different trials indicated that the best method was to use a small

amount of pulverised soil (1 g) and digest with nitric (AR) and perchloric acid (70 %

aristar) (1 :4) in 1 8 by 200 nim test tubes, in a block digester.

2.4.2 Acid Extraction Nitric/Perchloric (1:4) Method

All glass-wear was washed with high purity water and detergent (decon) before being

placed in a detergent bath (2% decon) for a several hours. They were then washed three

times with high purity water, soaked overnight in 10% nitric acid, rinsed thoroughly

with high purity water (Pennutit), dried in an oven.

An homogenous pulverised sample ( 1 g) was weighed into clean 200 mL test tubes (in

triplicate). One mL nitric acid (AR) was added to the test tube. The tubes were covered

with plastic film and the mixture was allowed to stand overnight in a block digester at

room temperature. The following morning, 4 mL perchloric acid (Aristar grade) was

added to the mixture. Gradually, over a period of I hour, the temperature was increased

83

to a maximum of 180' C and maintained at 180' C for 3 hours. Triplicate blank and

2 reference samples (in duplicate) were carried out with each batch of 50 samples;

The digest was allowed to cool before bringing to volume (20mL) with high purity

water (permutit). The digest was mixed thoroughly using a vortex mixer before being

transferred into SOmL polypropylene centrifuge tubes and centrifuged for 10 minutes at

12,000 rpm in a Beckman model N' J2-21MIE centrifuge.

The centrifuged samples were filtered through Whatrnan 541 filter paper into 50 mL polyethylene bottles and stored in a fridge at 4'C prior to analysis.

2.4.3 Atomic Absorption Spectrophotometer Analysis Procedures

Prior to being analysed for the vanous elements, the sample solutions (stored in

polyethylene bottles at 4•c) were allowed to reach room temperature and sonicated.

A number of elements (cobalt, copper, iron, manganese and zinc) were read directly or

diluted with high pwity water if necessary. Aluminium, calcium, magnesium, potassium

and sodium required the addition of releasing or ionisation agent to avoid chemical

interference. To avoid the necessity for separate dilutions128, lanthanum chloride (LaCI3)

(releasing agent) and caesium chloride (CsCl) (ionisation agent) were added to the

solutions. Aluminium and calcium were analysed using the nitrous oxide/acetylene

flame.

A stock solution containing 25,000 mg/L LaCI3 and 10,000 mg/L CsCI, in 2 M

hydrochloric acid, was prepared to give a final concentration of 5,000 mg!L lanthanum

and 2,000 mg/L caesium for the measurement of aluminium, calcium, magnesium,

sodium and potassium.

The blanks and the standards were prepared using the same proportions of reagents.

Perchloric acid was added to the standards to match the final acid concentration of the

84

sample solutions (10 % for copper, cobalt, iron, manganese and zinc and 1 % for the

other elements). The standards were prepared using BDH stock standard solutions (I 000

mg/L).

Prior to each set of measurements, the AAS was allowed to "warm up" for 20 min

minimwn before calibration. Three consecutive_ readings, each of 3·5 seconds

integration time, were obtained. Every I 0 samples, the instrument was recalibrated or

one standard was re-read. During the analyses, I M nitric acid followed by high purity

water were nebulised through the AAS instrument to keep the burner assembly as clean

as possible. The instrumental settings are given in Table 2.9.

TABLE 2.9 AAS instrument parameters

Elements Wavelength Slit Lamp current Background

(nm) (nm) (rnA) correction

AI 396.1 0.5 10 no

Ca 422.7 0.5 7 no

Co 240.7 0.2 7 yes

Cu 324.8 0.5 4 no

Fe 386.0 0.2 5 no

K 769.9 1.0 10 no

Mg 202.5 1.0 7 yes

Mn 279.5 0.2 5 no

Na 589.0 0.5 7 no

Zn 213.9 1.0 5 yes

Flame Standards Comment

(mg!L)

N20/C2H2 25.0 . 400 La/Cs

N20/C2H2 1.25 • 10 La/Cs

air-acetylene 0.3 13 - 5.0

air-acetylene 0.156 - 2.5

air-acetylene 25.0 - 200

air-acetylene 3.13 - 50 La/Cs



air-acetylene 0.625 - 5.0 La/Cs


2.4.4 Quality Assurance and Quality Control for the Analyses

85

The quality assurance and the quality control of the results were checked by using

reference materials and external laboratory analyses. As no termite mound reference

materials are available, BCSS�l and l\1ESS-l sediment reference material were selected

together with a soil reference material IAEA Soil 5. Four termite mound and three soil

samples were sent to an external laboratory and analysed by XRF and ICP-AES or ICP

MS. The four mound samples were used as internal reference samples. The precision

of the method was established by running 3 times (3-5 replica) the reference materials

(BCSS-1, MESS-I and IAEA SoilS) and the internal samples (A?l :Amitermes vitiosus

(Daly River), A53: Amitermes vitiosus (Elliott), Nl58: Nasutitermes triodiae (Daly

River), T99: Tumulitermes pastinator (Daly River), SA65 (Elliott soil), SN198 (Daly

River soil) and SN217 (Howard Springs soil).

During the analyses, the internal reference sample A?l was run with every batch of

samples. Another internal reference sample was run matching the type of termite

samples being read.

2.5 Infusion (Hot Water) Extractable Minerals

The "water infusion" of samples was prepared according to the description of the method

used by the Aboriginal community at Elliott (see chapter 1 . 1 .2.2c). For analytical

purposes, the Elliott water (bore) was substituted by distilled water. A total of 1 3

mounds (Amitermes vitiosus) (top and bottom sections) and 2 soils were analysed.

2.5.1 Extraction Process

An approximate 250g piece of Amitermes vitiosus mound was weighed and burned in

a fire, similar to the fire made by the Aboriginals in Elliott, for 15-20 minutes (until the

outside becomes black). The hot piece of mound was removed from the fire and placed

immediately in a 2L beaker containing 500 mL of high purity water (23°C). The weight

86

of the termite piece was rechecked as soon as the hot termitaria was in the water, by

difference between the total weight and the weight of the 2L beaker and the 500mL of

water. This was necessary as some material from the termite mound piece are destroyed

by the fire. The termitaria was left to infuse in the water for 1 0 minutes. After the first

5 minutes, the total weight (water + termite material) was determined again to calculate

the remaining amount of water, as some water evaporated on contact with the hot

termitaria. The preparation was then mixed and crushed until all big lumps of termitaria

were reduced to a sandy texture. The water extract was centrifuged in 50 mL

polypropylene centrifuge tubes for 15 minutes at 10,000 rpm in a Beckman model N"

J2-21MIE centrifuge, filtered through Whatman 541 filter paper and IOOmL of the

solution obtained was poured into a polyethylene bottle containing lmL of cone. nitric

acid. The soil samples (<2 mm fraction) being very sandy could not be burned in the

fire and were poured directly into hot water (85°)C; this was the temperature reached

by the mound mixtures during infusion. The determination of the element

concentrations (aluminium, calcium, cobalt, copper, iron, magnesium, manganese,

potassium, sodium and zinc) was performed using a Varian SpectrAA 40 Atomic

Absorption Spectrophotometer (AAS).


The analysis procedures were similar to those described previously in chapter 2.4.3. All

the standards were made I % in nitric acid to match the "infusion11 acidity. The AAS

instrumental settings are given in Table 2.10.

87

TABLE 2.10 AAS instrument parameters for analysis of hot water digest

Elements Wavelength Slit Lamp Background (nm) (nm) current correction

(mA)

A1 309.3 0.5 10 no Ca 422.7 0.5 10 no Co 240.7 0.2 7 yes

Cu 324.8 0.5 4 no

Fe 248.3 0.2 5 yes

K 769.9 1.0 1 0 no

Mg 202.5 1 .0 7 yes

Mn 279.5 0.2 5 no

Na 589.0 0.5 7 no

Zn 213.9 1.0 5 yes

2.6 Bioavailability of Fe in Termitaria

Flame Standards Comment (mg!L)

NP/C2H2 6.3 - 100 La/Cs

NP/C2H2 10 - 100 La/Cs

air�acetylene 0.313 - 5.0






air-acetylene 0.313 - 5.0 La/Cs


The bioavailable analysis ("in vitro11 analyses) method was a modification of the method

described by Narasinga Rao and Prabhavathi (1978)'22 for the determination of the

bioavailability of iron from foods with the addition of potassium fluoride to complex the

Fe(III)90 . The elements analysed were extended to the 9 other elements studied in this

project. The determination of the element concentrations was performed using a Perkin

Elmer Plasma 400 Inductively Coupled Plasma Argon Emission Spectrophotometer ICP

AES.

2.6.1 Bioavailability Extraction Trials

Different percentages of pepsin (0%, 0.1%, 0.5% and 5% w/v) were used in the

extraction procedure with samples A71 and Nl60, 3 times in triplicates. In view of the

results obtained, a 0.5 % w/v solution of pepsin was used for all extractions. The list

of sample analysed is given in Table 2 . 1 1 . The analyses were carried in triplicate on

a minimum of 3 mounds for each species and 2 soil ·samples at each site.

88

TABLE 2.1l Bioavailable test sample list

Site

Site I : Daly River

Site 2: Daly River

Site 3: Daly River

Site 4: Daly River

Site 5: Elliott

Sample number*

Tp0201, Tpl60!, Tp230!, ThO!dl, Th020!, Th030!, 050!, 0701

Av3 1D2, Av34D2, Av39D2, 0902, 1002

Tp34D3, Tp3603, Tp3803, Nt!OD3, Nt!203, Ntl403, 1703, 2203

Av4504, Av5004, Av6!D4, Nt2504, Nt2604, Nt2804, Nt3!D4,

Nt3404, Nt5604, Nt5904, Nt6504, Nt6804,Nt7204, 2304, 2604

AvO IE, Avl5E, Av22E, OlE, 02E

Site 6: Howard Springs Nt92H, Nt99H, Ntl06H, Tp45H, Tp60, Tp64, 27H, 29H

#: sample number explanation: section 2.1.3

2.6.2 Extraction Procedure

The extraction of soluble and ionisable iron was performed as follows:

Approximately 2.5g of termitaria and soil samples were incubated in a lOOmL conical

flask containing 50mL (0.5% w/v pepsin in 0.1 N HCl) at pH 1.35, at 37°C for 90

minutes in a metabolic shaker (200 rpm}. The pepsin used was from porcine stomach

mucosa (Sigma I :60,000; 2450 units/mg solid). After the incubation, the digests were

centrifuged for 1 5 minutes at 12,000 rpm in a Beckman J2-21M/E centrifuge and the

supernatant filtered through Whatman 541 filter paper into polyethylene bottles. An

aliquot of the filtrate (pH 1.35) was neutralised, in a 50mL polypropylene centrifuge

tube, with O. l N NaOH and the pH was adjusted to pH 7.5. The solution obtained was

centrifuged for 1 5 minutes at 12,000 rpm and filtered into 50 mL polyethylene bottles.

89


2.6.3.1 Total Concentrations

The total soluble iron, together with the elements AI, Ca, Co, Cu, K, Mg, Mn, Na, Zn

was determined on the pH 1.35 and pH 7.5 filtrates by ICP-AES. Prior the analyses,

the appropriated wavelength for each element was selected according to the sample

mineral composition in order to minimise the possible interference and to obtain a

maximum sensitivity. For the sodium values, the interference due to iron present in each

sample was calculated and deduced from the total sodium. The instrumental settings of

the ICP-AES are given in Table 2.12. The sodium was not measured at pH 7.5, due to

use of NaOH to neutralise pH 1.35 extract.

TABLE 2.12 ICP-AES instrument parameters (Pepsin-HCI extraction) (pH 1.35 and pH 7.5)

Elements Wavelength Standards Elements Wavelength Standards

(nm) (ppm) (nm) (ppm)

AI 396.152 0.500 - 50.00 K 769.490 0.500 - 50.00

Ca 3 1 7.933 5.000 - 50.00 Mg 279.553 5.000 - 50.00

Co 228.616 0.100 - 1.000 Mn 257.610 0 .I 00 - 1 .000

Cu 324.754 0.100 - 1 .000 Na 589.592 1 .000 - 10.00

Fe 238.204 0.100 - 0.500 Zn 213.856 0.100 - 1.000

0.500 - 50.00

90

2.6.3.2 Analysis of Fe(II)

Prior to being analysed for the ionisable iron (Fe(II)), the sample solution was first

acidified with {6M) HCI, then the Fe(III) was complexed as (FeF6l by addition of

potassium fluoride (2M) to the sample solution, l 0 minutes before adding a-a'

bipyridine (0.25 % w/v) and ammonium acetate · acetic acid buffer (pH 4.5). The Fe{II)

was measured colorimetrically in pH 1.35 and pH 7.5 extracts at the absorbance of 523

run on a Perkin-Elmer 552 UV-Visible Spectrophotometer, using a lcm plastic cell, as

HF (etching acid) is present in the filtrate. The calibration curve was established using

a range of standards from 0.05 ppm iron(ll) to I 0 ppm.


The precision of the technique was established in the same way as for the

nitric/perchloric analyses. A termitaria reference sample Nl60 (< 2 nun fraction) was

chosen and subjected to the analysis 3 times in triplicates. Sample N160 was also run

with every batch of samples.

CHAPTER lliREE

RESULTS

3 RESULTS

3.1 Site and Mound Characterisations

91

Four species of termite mounds (Plates 9-16), including the three species that are used

by the Aboriginal communities of Daly River or Elliott, have been studied from 4

geographically different localities as described in section 2.1.2.

3.1.1 Site Characterisations

A typical site with Nasutitermes triodiae and Amitermes vitiosus mounds in Daly River

is shown in Plate 9. The type of vegetation is open woodland (site 4) to woodland

(Eucalyptus) in Daly River, Howard Springs and Bertimah and open grassland with

scattered trees in Elliott.

3.1.2 Termite Species and Mound Characterisations

Four termite species were collected and identified. The identification was based on the

size and physical structure of the soldier caste, the termite distribution and the type of

mound. The identifications were later confirmed by Leigh Miller from CSIRO Division

of Entomology, Canberra.

All the termites studied belong to the recent family: Termitidae (termites with worker

caste) and are grass-eating termites. Three belong to the sub-family Nasutitennitinae:

Nasutitermes triodiae (Froggatt) (Plates 10 and I I), Tumulitermes pastinator (Hill)

(Plates 12 and 13) and Tumulitermes hastilis (Froggatt) (Plate 14). They have nasute

head (Figure 3.1). Nasutitermes sp are distinctive with the soldier's head is not

constricted (Figure 3.1-A), while Tumulitermes sp have the soldier's head constricted

near its centre (Figure 3.1-B). Amitermes vitiosus Hill (Plates I S and 16) belongs to the

~

,....

>

--'

-rl

-c 3 0 :::

::l c.. .,.

~ ~ ~ ::- " " "'0 et " ~

l6

93

Amitermitinae subfamily. Amitermes sp have mandibulate heads with mandibles toothed

and sabre-shaped (Figure 3 . I -C).

The termitaria have characteristic features, although there is a degree of variation in

mound shape within species. A general representation of the size (height +

circumference) of the termitaria sampled at each site, is given in Figure 3.2. As seen

in Figure 3.2, Amitermes vitiosus and Tumulitermes hastilis have mounds that are small

and usually narrow, those of Tumulitermes pastinator are low but wide and those of

Nasutitermes triodiae are tall and large.

A. Nasutitermes sp

Nasute head

B. Tumulitermes sp

Mandibulate head

C. Amitermes sp

FIGURE 3.1 Dorsal view ofnasute head: A: Nasutitermes sp and B: Tumu/itermes sp and mandibulate head: C : Amitermes sp. (from Hadlington, 198764)

PLATE 10 Nasutitermes triodiae mound (3m height), Daly River, site 3 .

PLATE I I Vertical section of Nasutitermes triodiae mound at site 3, showing compact basal portion-galleries and nursery (middle part of the mound, ground level).

\C .a:..

1000

BOO

600 8

400

200

0

FIGURE 3.2

3.1.2.1

95

Av Nt

8 CIRCUM [l HEIGHT

z 4 5 . , 3 • 3 • • 7

SITE

Size comparison (height + circumference in em) of Amitermes vitiosus (Av), Tumulitermespastinator (Tp1 Nasutitermes triodiae (Nt) and Tumulitermes hasti/is (Th) mounds at different sites.

Nasutitermes triodiae (Froggatt)

These nasute soldiers (4.5 ± 0.25 mm) have a dark brown head extended into a thin

nasus (Figure 3.1-A) through which they can expel a sticky repellent secretion associated

with the defence against ants and other enemies. Nasutitermes triodiae occur widely in

Northern Australia. They are also known as "spinifex termites". They construct various

types of mounds that are the largest of any of the Australian species, reaching a height

of 6 metres. The mounds collected in Daly River are the "cathedral" type, (Plates 9, 10

and I I), they are only found in the Top End, north of Pine Creek. Elsewhere,

Nasutitermes triodiae builds large mushroom-shaped mounds or columnar mounds

without the Top End elaborations151•72,

In these mounds, the inner region (beside the nursery N) is solid and extremely hard.

Around this solid central core, there is a zone of open galleries and cells in which

chaffed grass (8.61 ± 0.51 mm) is stored in considerable quantities during the dry

season97• As the mound grows, the outer storage chambers are abandoned and re-packed

with soil.

PLATE 1 2 Tumulitermes pastinator mound (70cm height). Daly

River, site 3. Vertical section of Tumulitermes pastinator mound

at site 3 (Daly River), showing alveolar type of

structure and the nursery (N).

\C �

97

3.1.2.2 Tumulitermes pastinator {Hill)

Tumulitermes pastinator is a small species of variable size and colour; the soldiers

measure 3.5 to 3.75 mm long, they have a nasute head usually light to dark brown72•

The species extends across Northern Australia from Queensland to Western Australia52•

The mounds are generally low dome-shaped structures (Plate 12) about 60-80 em high and 60-90 em in diameter, but occasionally they reach a height of 1.2 m and a basal

diameter of 1 .5 m. The outer wall is made of repacked soil material and is very thin and

dense. It covers an open alveolar interior made of softer repacked soil material (Plate

13). The vast number of chambers and galleries are packed with fragments of chaffed

grass stems. The central nursery [N] (Plate 13 ground level) is made of a soil/carton

mixture97

3.1.2.3 Amitermes vitiosus Hill

Amitermes vitiosus is a variable species with a dark orange head. The soldiers measure

4-5 mm and are of the mandibulate type72 (Figure 3.1-C). It is a fairly common species

in the northern parts of Queensland and the Northern TerritorY2• It is often found in

association with other grass�feeding mound�building species, Nasutitermes triodiae

(site 4) and Tumulitermes hastilis. The mounds of this termite are remarkable for their

abundance in certain localities and for their diversity in form. Their structures are

intensely hard (concrete� hard material), with a thin, undifferentiated outer wall which

is often deeply sculptured (Plates 14 and 15). They range in colour from light gray

(site 2) to dark gray (site 4) and to deep mahogany red (site 5, Plates 14 and 15)

according to the colour of the surrounding soi1'2• The commonest form consists of a

colunmar mound up to 1.2 m high with a basal diameter up to 60 em (site 2). On sandy

soil, it builds small conical mounds (site 5). On areas subject to seasonal flooding (such

as site 4), the mounds closely resemble those of Amitermes meridionalis in being

laterally flattened and oriented more or less on a north·south axis (Plate 9).

PLATE 14 Amitermes vitiosus mound (50cm height), Elliott. PLATE IS Vertical section of Amitermes vitiosus mound in

Elliott showing the concrete hard structure.

\C 00

99

The mounds show little obvious differentiation in gross structure. The internal structure

consists of repacked soil with interconnected galleries. The gaJleries are usually lined

by very fine, black organic residues (Sites 2 and 4). In the outer part of the mound,

they are often filled with fragments of grass, leaves and organic materials, including the

bodies of dead termites 38.

3.1.2.4 Tumuliterme.� hastilis (Frogatt)

The soldiers measure 3.5-4.5 mm. The species has a wide distribution in the inland low

rainfall areas of Queensland, Northern Territory, Western Australia and South Australia.

It builds tall narrow mounds up to 1 . 5 m height and 50 em wide at ground level52 In

Daly River site I, the mounds are relatively small and narrow (see Plate 16), they do not

exceed 90 em height. The construction material is mostly soil97. The interior consists

of large numbers of small chambers stored with grass ·and pieces of unidentifiable

vegetable debris.

Plate 16 Tumulitermes hastilis mound (55cm height), Daly River. site I .

� <:> TABLE 3.1 Quality control of selected elemental composition of reference material (BCSS-1, MESS-I <:>

and 1AEA SOILS), following perchloric/nitric acid (4:1) extraction (mgl100g).

Element Reference material

BCSS-1 MESS-I IAEA SOILS

Certified HNO/HCI04 Certified HNO/HCI04 Certified HNO,IHCIO,

values n=lO N"alues n=9 values n=l l

Aluminium 6260 ± 217 4245 ± 372 5837 ± 201 3140 ± 80 8190 ± 280 4333 ± !57

Calcium 543 ± 53 412 ± 14 482 ± 46 261 ± 7 2200 *nc 1341 ± 56

Cobalt 1 . 14 ± 0.21 0.98 ± 0.05 1.08 ± 0.19 0.93 ± 0.06 1.48 ± 0.08 1.08 ± 0.04

Copper 1.85 ± 0.27 1.49 ± 0.05 2.51 ± 0.38 2.22 ± 0.09 7.71 ± 0.47 7.21 ± 0.43

Iron 3287 ± 98 3539 ± 201 3050 ± 175 2921 ± 189 nd 4472 ± 192

Potassium 1801 ± 33 1088 ± 175 1859 ± 33 775 ± 78 1860 ± ISO 855 ± 123

Magnesium 1471 ± 139 1321 ± 80 868 ± 54 697 ± 40 1500 *nc 1002 ± 61

Manganese 22.9 ± 1.5 20.4 ± 0.5 5 1 .3 ± 2.5 40.2 ± 1.4 nd 70.5 ± 1.4

Sodium 2018 ± 156 959 ± 65 1855 ± I l l 690 ± 58 1920 ± 1 1 0 93.5 ± 9.3

Zinc 1 1 .9 ± 1.2 10.1 ± 0.2 19.1 ± 1.7 17.2 ± 0.9 36.8 ± 0.8 36.0 ± 2.1

nd: not detennined • nc: not certified, OSS (Jabiru) value

101

3.2 Acid Extractable (PerchloridNitric Acids) Selected Elements from Termite

Mounds and Soil Together with Particle Sizes.


The results of the quality control and quality assurance are shown in Tables 3.1, 3.2 and

3.3. Table 3 .1 shows the results of selected elemental composition of reference material

(BCSS-1, MESS-I and IAEA SOILS) following perchloric/nitric acids (4:1) extraction.

This extraction does not result in a total concentration in all elements. However, the

efficiency for Fe is greater than 95 % for all the reference materials and the extraction

efficiencies for Co, Cu, Mg, Mn and Zn are generally greater than 80 %. The Ca

extraction efficiency varies between reference materials (54 % in MESS-I to .76 % in

BCSS-1). Na and K are poorly extracted from soils and sediments. The efficiency

being of the order of 42 to 60 % for K and 5 to 48 % for Na.

. The extraction efficiency depends on the mineralogical nature of the sample. It was

therefore important to check with reference material more closely matched to the

samples of this study. As there is no termitaria reference material available, internal

reference materials were established. Four termite mound samples: Amitermes vitiosus

(Daly River, site 4), Amitermes vitiosus (Elliott, site 5), Tumu/itermes pastinator (Daly

River, site 1), Nasutitermes triodiae (Daly River, site 4) and three soil samples: Daly

River (site 4), Elliott (site 5) and Howard Springs (site 6) were sent to a private

laboratory for comparative analyses. The termitaria samples were analysed by X-ray

fluorescence for AI, Ca. Fe, K, Mg and Zn and following a mixed acid digestion (HCI,

HCI04 and HF) by Inductively Coupled Plasma Optical Emission Spectrometry (ICP

OES) for all the selected elements but Co and by Inductively Coupled Plasma Mass

Spectrometer (ICP-MS) for Co. Soils were analysed by ICP-OES following mixed acid

digestion (HCI, HCI04 and HF)

Tables 3.2 and 3.3 show the results of the selected elemental composition of the internal

reference materials (mounds and soils) following perchloric/nitric acids ( 4: 1) extraction.

TABLE 3.2 Quality control of selected elemental composition of internal reference termitaria material (Av44D4, Av22E, Tp23Dl and Nt24D4), following perchloric/nitric acid (4:1) extraction (mg/lOOg).

Element Internal reference tennitaria material®

Av44D4 Av22E

Ext.Lab HNO/HCI04 Ext. Lab HNO/HC104 -

XRF lCPj n=55 XRF lCPt n=25

Aluminium 4567 4800 3986 ± 143 2911 3420 3614 ± 151 .

Calcium 93 109 93 ± 3 122 151 137 ± 4

Cobalt 0 #0.54 0.42 ± 0.02 - #0.43 0.31 ± 0.03

Copper - 1.10 0.91 ± 0.03 - 1.00 0.83 ± 0.03

Iron 1350 1330 1271 ± 26 1476 1580 1625 ± 87

Potassium 1 1 12 1 170 702 ± 68 158 198 161 ± 15

Magnesium 15 1 159 139 ± 5 30 92 96 ± 5

Manganese 7.80 10.3 7.34 ± 0.22 7.80 9.90 7.55 ± 0.32

Sodium - 52.1 30.0 ± 2.4 - 10.7 5.7 ± 0.3

Zinc - 0.80 0.52 ± 0.03 1 .30 1 .00 ± 0.05

@ : for internal reference termitaria material number explanations see chapter 2J.3 Abbreviations: Ext.Lab = external laboratory; XRF = X-Ray Fluorescence; ICP - Inductively Coupled PLasma t: ICP mixed acid digestion = JICI, JICIO,, fiF; II: ICPMS - Inductively Coupled Plama Speclrophotometer

Tp23Dl

Ext. Lab HNO/HC104

XRF lCPt n=32

3514 3590 3194 ± 1 1 8

21.4 34.0 26.5 ± 1.0

- #0.37 0.29 ± 0.03

- 0.80 0.58 ± 0.02

1336 1330 1354 ± 52

780 781 526 ± 45

42 78 68 ± 4

7.80 5.00 3.16 ± 0.08

- 46.0 31 .4 ± 2.5

- 0.70 0.43 ± 0.03

Nt2404

Ext. Lab HNO/HC104

XRF lCPt n=33

4869 5250 4608 ± 2 1 1

21.4 37.5 28.8 ± 1.3

- #0.39 0.29 ± 0.03

- 1.00 0.89 ± 0.03

2000 2180 2149 ± 98

913 1020 652 ± 69

54 1 14 100 ± 4

7.80 3.70 2.44 ± 0.12

- 39.0 22.1 ± 2.8

- 0.60 0.43 ± 0.04

� <:> N

103

The method shows good extraction efficiencies compared to the XRF external laboratory

results, except for Mn in the Tumulitermes pastinator and Nasutitermes triodiae mound

samples; and good recoveries compared to ICP-OES and JCP-MS, except for Na (where

percentage recovery varies from 53 to 68 % in the mounds and 43 to 63 % in the soils).

The K percentage recoveries were highest in the Elliott samples: 100 % in the mound

and 86 % in the soil, while at the other sites it was arotu1d 60 to 70 %.

The general precision of the method is indicated by the selected element standard

deviations given in Tables 3.1, 3.2 and 3.3. These values are the total of the values

obtained for all the runs, during testings and analyses. They were usually very low

(below 5 %) but K and Na were higher (± 10 %).

3.2.2 Overview, General Correlation

A global overview of termite mound selected elements and particle sizes data from

different areas and different species is presented in Table 3.4. It shows strong negative

and positive correlations between most of the variables. Out of the 91 variable

cOrrelations, 72 were significantly correlated. For example, the iron was positively

correlated to aluminium, cobalt, copper, manganese, zinc, clay and fine sand. It was

negatively correlated to potassium, magnesium, sodium, silt and coarse sand. No

significant correlation was observed between iron, calcium and silt. The calcium was

positively correlated to copper, manganese and zinc and negatively correlated to

potassium and sodium. The clay was significantly correlated to most of the variables:

positively correlated to aluminium, cobalt, copper and iron and negatively correlated to

potassium, magnesium, silt and coarse sand. No significant correlation has been found

between clay, calcium and fine sand.

While a complete study of the correlations would be interesting, it is not the focus of

the project which has concentrated on Aboriginal community use ofterrn.itaria. For this

purpose, a finer detailed investigation between: age of sample collection, sample

position, size of mound, mounds of different species and different sites has been

necessary.

� = ...

TABLE 3.3 Quality control of selected elemental composition of internal reference soil material (OlE, 25D4 and 29H), following perchloric/nitric acid (4:1) extraction (mg/IOOg).

Internal reference soil materia!®

OlE 2504 29H

Element Ext. Lab HNO/HC104 Ext. Lab HNOiHC104 Ext.Lab HNO/HC104 ICPt n=15 ICPt n=15 ICPt n=IS

Aluminium !610 1677 ± 48 2600 2216 ± 159 3340 3587 ± 76

Calcium 55.1 46.2 ± 1.4 13.5 6.8 ± 0.5 59.8 51.4 ± 1 5 Cobalt #0.32 0.21 ± 0.02 #0.33 0.20 ± 0.03 # 0.92 0.70 ± 0.02

Copper 0.70 0.55 ± 0.01 0.60 0.48 ± 0.02 1.50 1.22 ± 0.03

Iron ! 100 1167 ± 42 820 844 ± 20 7040 7648 ± 209

Potassium ! 18 101 ± 6 746 449 ± 62 50 28 ± 2

Magnesium 42 44 ± 3 56 49 ± s 3 1 35 ± 3

Manganese 7.50 5.66 ± 0.13 3.90 2.31 ± 0.09 14.9 12.1 ± 0.34

Sodium 9.80 4.25 ± 0.74 32.8 20.7 ± 3.80 7.90 4.39 ± 0.55

Zinc 0.70 0.51 ± 0.02 0.50 0.25 ± 0.02 2.60 2.60 ± 0.10

@: for C1(planation of soil reference material number n:fi:r to chapter 2.3.1 t: mixed acid digestion - HCI, HCIO,, HF; #: ICPMS

TABLE 3.4 Pearson correlation (PC) matrix and probabilities (P t) of selected elements and particle size of 87 termite mounds (n=189) of all the species and sites studied (depth=l).

Element/ Aluminium Calcium Cobalt Copper Iron Potassium Magnesium Particle size PC p PC p PC p PC p PC p PC p PC p Aluminium 1.000 ...

Calcium 0.051 N 1.000 ...

Cobalt 0.749 ••• 0.056 N 1.000 •••

Copper 0.801 ... 0.160 • 0.818 ••• 1.000 ••• Iron 0.842 ... 0.037 N 0.764 ••• 0.817 ••• 1.000 •••

Potassium -0.465 ... -0.439 ••• -0.505 ••• -0.536 ••• -0.561 ••• 1.000 •••

Magnesium -0.182 • -0.061 N -0.222 •• -0.172 • -0.163 • 0.675 ••• 1 .000 ••• Manganese 0.146 • 0.504 ••• 0.329 ... 0.322 ••• 0.371 ••• -0.242 •• 0.345 •••

Sodium -0.404 ... -0.350 ••• -0.483 ••• -0.550 ••• -0.605 •• • 0.800 ••• 0.262 •••

Zinc 0.468 ... 0.246 •• 0.5 1 1 ••• 0.670 ••• 0.661 ••• -0.454 ••• .0219 •• Clay 0.789 ... -0.004 N 0.553 ••• 0.635 ... 0.618 ••• -0.322 ••• -0.153 • Silt -0.215 .. 0.083 N -0.100 N -0.105 N -0.121 N 0.395 ••• 0.718 ••• F.sand 0.254 ... -0.000 N 0.362 ••• 0.312 ••• 0.314 ••• -0.357 ••• -0.478 ••• Coarse sand -0.667 ... -0.013 N -0.636 ••• -0.639 ••• -0.625 ••• 0.258 ••• 0.086 N

Element/ Manganese Sodium Zinc Clay Silt Fine sand Coarse sand Particle size PC p PC p PC p PC p PC p PC p PC p Manganese 1.000 ...

Sodium -0.480 ... 1.000 ••• Zinc 0.648 ... -0.722 ••• 1.000 •••

Clay -0.080 N 0.267 ••• 0.294 ••• 1.000 ••• Silt 0.489 ... 0.074 N 0.320 ••• -0.459 ••• 1.000 ••• Fine sand -0.041 N -0.080 N 0.044 N -0.044 N -0.076 N 1.000 ••• Coarse sand -0.156 • 0.141 N -0.387 ... -0.433 ••• -0.151 • -0.714 ••• l .OOO ***

f: N: P>O.O$; *: O.Ol<P<O.O:>; *": O.OOI<P<O.Ol; •u: P<O.OOI � = "'

106

3.2.3 Detailed Mound Study

One mound of each termite species selected by the Aboriginal communities of Elliott

and Daly River, was analysed in detail. The detailed study included comparisons

between:

a) the age of the outside material of the moWld (old and new) in Nasutitermes triodiae

and Tumulitermes pastinator mounds;

b) the depth of sample collect where: depth�O, represents the Q.J em fraction of the

outside mound; depth= 1 , represents the 0� 10 em fraction of the outside mound; and

depth=2, represents the 0-10 em taken from the inside central axis of the mound.

Samples were taken from depths= 1 and 2 for Amitermes vitiosus and from depths=O,

I and 2 for Tumulitermes pastinator and Nasutitermes triodiae mounds;

c) the vertical position of the sample in the mound: top, middle and bottom

The physical characteristics of the termitaria selected are given in Table 2.2 and the

detailed mound sample summary for the 3 termitaria is given in Table 2.3. The detailed

results of the analyses are given in Appendices lc, le, lg, lllc, llle, lllg.

3.2.3.1 Amitermes vitiosus (Elliott, Site 5)

The effects of depth and position on the selected elements and particle sizes of the

mound together with ANOV A probability of differences are presented in Table 3.5 and

Table 3.6 respectively.

107

TABLE 3.5 Amitennes vitiosus mound detailed study (Elliott, site 5): depth effects on selected elements (mg/lOOg) and particle sizes (o/o) (mean ± standard deviation) together with ANOV A probability of differences (P) between depths.

Element/ Depth-I Depth-2 p Particle size n""6 n=3 � Aluminium 3428 ± 128 3548 ± 191 N

Calcium 136 ± 17 127 ± 22 N

Cobalt 0.32 ± 0,03 0.35 ± O.Ql N

Copper 0.91 ± 0.03 0.92 ± 0.03 N

Iron 1644 ± 43 1666 ± 24 N

Potassium 154 ± 2.8 160 ± 3.0 N

Magnesium 92 ± 2.1 92 ± 5.5 N

Manganese 7.93 ± 0.33 7.76 ± 0.23 N

Sodium 6.19 ± 0.32 5.98 ± 0.43 N

Zinc 1.09 ± 0.10 1.10 ± 0.11 N

Clay 16.5 ± 6.5 20.8 ± 2.3 N

Silt 10.4 ± 5.0 9.7 ± 5.3 N

Fine sand 36.3 ± 6.0 33.8 ± 1.1 N

Coarse sand 38.0 ± 1.3 37.5 ± 0.9 N l: N: P>O.OS

In the mound studied, there were no statistically significant differences associated with

depth of the sample in the mound.

The ANOV A shows that there are highly significant differences in calcium, magnesium

and zinc with position (Table 3.6). The pairwise comparisons (Tukey test) indicate that

the differences are associated with increases in concentrations in the top and middle

sections of the mound (Figure 3.3).

108

160 l 1.2 -

� Bottom 0 Middle

120 1 Fj l Lc Too_l 09 0, 0 0 -' 0 80 � El � El � 06 5 \1 0 0 40 � El � El � 03

0 -'--- 0.0 I 1=1 ,

FIGURE 3.3

c. MINERAL

Mg zo MINERAL

Position effects (mean ± SE) on calcium, magnesium and zinc (mg/lOOg) in Amitermes vitiosus mound sampled in Elliott (site 5) at depths I and 2

109

TABLE 3.6 Amitermes vitiosus mound detailed study (Elliott, site 5): position effects on selected elements (mg/lOOg) and particle sizes (%) (mean ± standard deviation) together with ANOV A probability of differences (P) between positions.

Element/ Top Middle Bottom Particle size n=3 n=4 n=2

Aluminium 3529 ± 135 3521 ± 1 1 7 3271 ± 82

Calcium 133 ± 3.4 147 ± 5.5 105 ± 1 .00

Cobalt 0.34 ± 0.03 0.32 ± 0.02 0.33 ± 0.04

Copper 0.90 ± 0.02 0.93 ± 0.03 0.92 ± 0.05

Iron 1655 ± 70 1653 ± 21 1641 ± 1.0

Potassium 158 ± 3.5 153 ± 3.4 160 ± 3.9

Magnesium 94 ± 2.3 93 ± 0.8 87 ± 1.3

Manganese 7.88 ± 0.40 7.93 ± 0.32 7.74 ± 0.15

Sodium 6.36 ± 0.21 6.16 ± 0.31 5.67 ± 0.01

Zinc 1 . 15 ± 0.63 1.14 ± 0.02 0.94 ± 0.06

Clay 12.1 ± 6.1 20.6 ± 3.1 21.3 ± 0.4

Silt 15.5 ± 2.4 8.4 ± 3.2 5.6 ± 0.7

Fine sand 39.9 ± 7.0 32.5 ± 0.9 34.9 ± 0.3

Coarse sand 37.3 ± 1 . 1 38.3 ± 1.2 37.6 ± 1.4

l: N: P>O.OS; . . : O.OOJ<P<O.OI

3.2.3.2 Tumulitermes pastinator (Daly River, Site 3)

p t N ..

N

N

N

N . .

N

N ••

N

N

N

N

The effects of age on the selected elements and particle sizes of the mound together with

the ANOVA probability of differences are presented in Table 3.7. The results of the

ANOV A show that the effects of age on the proportion of the selected elements and

particle sizes were not significant for the Tumulitermes pastinator mound at site 3

(P>0.05).

110

80

60 � ... a. 0 0 � ' § 40 � 1111! Sl 0 0

20 � .I.Ellli!

1000

750 � ' IIIII

500 � 1 rn

250 � 1 rn

1 2

g

6

3

l8l Depth-2 g Depth•1 D Depth .. O

0 -'-----'- 0 -'---__u o L--.t...r;:

FIGURE 3.4

FIGURE 3.5

c.

40 ,

I 30

� z

K MINERAL

Mo

Depth effects (mean ± SE) on calcium, potassium and manganese (mg!IOOg) in Tumulitermes pastinator mound sampled in Daly river (site 3)

l ll!l Deoth•2 El Depth-1 0 Depth-0

T

� 20 a: w a_

10

0 I I '7 Sill Coarse sand PARTICLE SIZE

Depth effects (mean ± SE) on silt and coarse sand (%) in Tumulitermes pastinator mound sampled in Daly river (site 3)

111

TABLE 3. 7 Tumulitermes pastinator mound detailed study: age effects on selected elements (mg/lOOg) and particle sizes (o/o) (mean ± standard deviation) together with ANOVA probability of differences (P) between ages. (Deptb=O).

Element/ Old material New material p Particle size n=5 n=2 + Aluminium 3983 ± 498 3986 ± 161 N

Calcium 16.5 ± 1.3 24.1 ± 7.6 N

Cobalt 0.44 ± 0.04 0.45 ± 0.01 N

Copper 0.86 ± 0.08 0.85 ± 0.01 N

Iron 2942 ± 250 2912 ± 108 N

Potassium 701 ± 83 755 ± 67 N

Magnesium !55 ± 10 164 ± 24 N

Manganese 5.43 ± 0.41 5.96 ± 0.94 N

Sodium 14.9 ± 3.0 15.3 ± 1.5 N

Zinc 1 .24 ± 0.10 1 .26 ± 0.14 N

Clay 18.3 ± 1.3 19.4 ± 0.3 N

Silt 16.4 ± 1.0 16.9 ± 0.1 N

Fine sand 37.0 ± 0.3 38.4 ± 4.1 N

Coarse sand 31.5 ± 1.7 30.2 ± 5.0 N

I' N: P>O.OS

The effects of depth and position on the selected elements and particle sizes of the

Tumulitermes pastinator mound together with the ANOV A probability of differences are

presented in Table 3.8.

The ANOVA shows a highly significant difference (P<O.Ol) in calcium, potassium,

manganese, silt and coarse sand and a significant difference in clay (P<0.05) associated

with depth. The pair-wise comparison between depths indicates that the inner section

of the mound ( depth=2) contains a higher level of calcium, potassium, manganese

(Figure 3.4), higher silt aod lower coarse saod particles (Figure 3.5).

� � TABLE 3.8 Tumulitermes pastinator mound (Daly River, site 3) detailed study: depth and position effects on selected

...

elements (mg/lOOg) and particle size (%) (mean ± standard deviation) together with ANOVA probability of differences (P) between depths and positions (at deptb=l).

Element/ Depth=O Depth= 1 Depth=2 P (Depths) Top Middle Bottom P (Positions)

Particle size n=7

Aluminium 3984 ± 412

Calcium 18.7 ± 4.9

Cobalt 0.44 ± o.m Copper 0.85 ± 0.08

Iron 2934 ± 210

Potassium 717 ± 78

Magnesium 158 ± 13

Manganese 5.58 ± 0.57

Sodium 15.0 ± 2.5

Zinc 1.24 ± 0.10

Clay 18.6 ± 1.2

Silt 16.5 ± 0.9

Fine sand 37.4 ± 1.8

Coarse sand 3 1 . 1 ± 2.6

n=6

3826 ± 446

23.3 ± 3.9

0.42 ± 0.04

1.50 ± 0.55

2917 ± 198

651 ± 79

152 ± 19

5.97 ± 0.83

13.8 ± 2.1

1.54 ± 0.28

2 1 .2 ± 2.4

15.6 ± 2.2

36.9 ± 2 . 1

28.8 ± 2.9

n=4

4039 ± 269

60.2 ± q.o 0.48 ± 0.04

1.34 ± 0.46

2939 ± 186

783 ± 61

175 ± 16

10.0 ± 2.1

16.8 ± 1.4

1.66 ± 0.31

20.2 ± 1.3

18.7 ± 0.8

38.1 ± 2.4

23.7 ± 2.7

I ' N: P>O . O S ; * : O . O l<P< O . O S ; * * : 0 . 00l<P<0.01

t n=2 n=2 n=2 t N 3795 ± 233 4280 ± 382 3403 ± 145 N

•• 23.8 ± 1.2 19.8 ± 1.71 26.2 ± 5.34 N

N 0.42 ± 0.04 0.44 ± 0.06 0.39 ± 0.01 N

N 1.22 ± 0.64 1.81 ± 0.25 1.48 ± 0.82 N

N 2821 ± 204 3117 ± 175 2814 ± 55 N

• • 629 ± 51 732 ± 87 593 ± 20 N

N 165 ± 3.2 159 ± 20.8 131 ± 5.6 N

• • 6.08 ± 0.77 5.32 ± 0.40 6.50 ± 1.10 N

N 1 3 . 1 ± 3.5 15.2 ± 1.80 13.0 ± 0.57 N

N 1.46 ± 0.28 1.75 ± O.Q2 1.43 ± 0.42 N

• 21.2 ± 1.9 23.1 ± 2.7 19.4 ± 2.3 N

• • 16.1 ± 1.2 17.4 ± 1.9 13.2 ± 0.3 N

N 35.9 ± 1.0 37.5 ± 0.1 37.4 ± 4.1 N

• • 28.8 ± 1.7 26.0 ± 0.8 31.7 ± 2.5 N

113

There are no significant differences associated with position of elements or particle sizes

content at depth= ! (Table 3.8). The other depths (0 and 2) were not investigated

because of the size of the mound.

3.2.3.3 Nasutitermes triodiae (Daly River, Site 3)

The effects of age on the selected elements and particle sizes of the mound together with

the ANOVA probability of differences are presented in Table 3.9. The results of the

ANOV A show that the effects of age on the proportion of the selected elements and particles size were not significant for the Nasutitennes triodiae mound at site 3 (P > 0.05).

Table 3.10 shows that at Daly River site 3, there were significant differences for cobalt,

iron, sodium and coarse sand with depth. The pair-wise comparison between depths

indicates that there was an increase in cobalt in the inner p;rrt of the mound (depth =2),

and an increase of iron, sodium and coarse sand in the outer part (depth =0) (Fignre 3. 6).

The calcium concentration was higher at depth=2, however the increase was not

significant due to a very high standard deviation.

At depth= I , the AN OVA shows significant differences in calcium and coarse sand with

positions (Table 3 . 1 0). The calcium concentration was higher at the bottom position of

the mound and the coarse sand was lower at the top (Figure 3.7).

114

3600

� 3500 a 8 -

' ..§' 3400

0 z 0 ()

3300

3200

,, MINERAl

FIGURE 3.6

30

" 8 25

r � 20

19 23 l!':l Qgpth-2

El Deoth- 1

0 Depth•O

18 22

17 � 21 w 0 �

16 � 20

15 19

14 18 No Coarse sand

MINERAL PARTICLE StZE

Depth effects (mean ± SE) on iron and sodium (mg/lOOg) together with coarse sand (%) in Nasutitermes triodiae mound sampled in Daly river (site 3)

22

2 1

� iii � 20

1 9

� Bottom

0 Middle

El Top

1 5 .L.._----=:1..,.- 18 .1._---=L,--

FIGURE 3.7

Co

MINERAL

Coarse sand

PARTICLE SIZE

Position effects (mean ± SE) on calcium (mg/JOOg) and coarse sand (%) in Nasutitermes triodiae mound sampled in Daly River (site 3)

115

TABLE 3.9 Nasutitermes triodiae mound detailed study: age effects on selected elements (mgllOOg) and particle sizes (%) (mean ± standard deviation) together with ANOV A probability of differences (P) between ages. (Depth=O).

Element/ Old material New material p Particle size n=l n=S t Aluminium 4772 ± 158 4722 ± 303 0.776 N

Calcium 23.1 ± 3 . 1 25.1 ± 4.7 0.501 N

Cobalt 0.52 ± 0.03 0.53 ± 0.02 0.718 N

Copper 0.82 ± 0.14 0.90 ± 0.19 0.490 N

Iron 3493 ± 169 3596 ± 232 0.485 N

Potassium 940 ± 81 987 ± 72 0.389 N

Magnesium 265 ± 37 252 ± 16 0.550 N

Manganese 9.61 ± 0.89 9.79 ± 1.79 0.859 N

Sodium 18.1 ± 1.9 18.4 ± 2.3 0.836 N

Zinc 1.63 ± O.o7 1 .62 ± 0.09 0.903 N

Clay 19.8 ± 1.6 23.6 ± 7.4 0.355 N

Silt 25.5 ± 5.0 26.0 ± 7.5 0.908 N

Fine sand 33.1 ± 1.8 33.3 ± 1.9 0.900 N

Coarse sand 23.5 ± 4.2 21.5 ± 1.7 0.350 N

t: N: P>O.OS

� TABLE 3.10 Nasutitennes triodiae mound (Daly River site 3) detailed study: depth and position effects on selected

� "' minerals (mg!IOOg) and particle size (%) (mean ± standard deviation) together with AN OVA probability of differences (P) between depths and positions (at depth=!).

Element/ Depth effects Position effects

Particle size Depth=O Depth=l Depth=2 P (Depths) Top Middle Bottom P (Positions) n=9 n=6 n=5 t n=2 n=2 n=2 t

Aluminium 4744 ± 236 4488 ± 167 4480 ± 234 N 4439 ± 26 4602 ± 502 4422 ± 312 0.610 N

Calcium 24.2 ± 3.9 23.1 ± 4.7 32.2 ± 23.6 N 19.9 ± 1.3 20.5 ± 2.6 28.8 ± 2.1 0.039 .

Cobalt 0.52 ± 0.02 0.52 ± 0.04 0.57 ± O.Q2 • 0.54 ± 0.04 0.52 ± 0.06 0.50 ± 0.01 0.633 N

Copper 0.87 ± 0.16 0.97 ± 0.30 0.98 ± 0.25 N 0.93 ± 0.37 0.98 ± 0.48 1.02 ± 0.26 0.970 N

Iron 3550 ± 202 3434 ± 124 3245 ± 139 • 3352 ± 120 3563 ± 108 3387 ± 1.0 0.195 N

Potassium 967 ± 76 897 ± 65 949 ± 95.5 N 862 ± [ [ 1 948 ± 12 881 ± 18 0.481 N

Magnesium 258 ± 26 256 ± 25 260 ± 22.5 N 239 ± 14 261 ± 8.3 268 ± 44 0.585 N

Manganese 9.71 ± 1.38 9.38 ± 0.99 9.69 ± 2.47 N 8.62 ± 0.91 9.84 ± 0.92 9.68 ± 1.21 0.515 N

Sodium 18.3 ± 2.0 16.5 ± 2.0 14.6 ± 1.3 • 14.7 ± 1.4 16.2 ± 0.1 18.4 ± 2.2 0.188 N

Zinc 1.63 ± 0.08 1.68 ± 0.18 1.74 ± 0.21 N 1.60 ± 0.07 1.74 ± 0.33 1.69 ± 0.15 0.803 N

Clay 21.9 ± 5.7 19.4 ± 1.0 21.9 ± 3.6 N 19.1 ± 1.0 19.3 ± 0.8 19.8 ± 1.6 0.833 N

Silt 25.8 ± 6.1 29.4 ± 1.3 29.8 ± 3.2 N 29.4 ± 1.4 30.5 ± 1.6 28.5 ± 0.4 0.391 N

Fine sand 33.2 ± 1.8 33.6 ± 2.1 32.9 ± 2.2 N 35.2 ± 1.0 31.4 ± 2.3 34.1 ± 0.6 0.164 N

Coarse sand 22.4 ± 3.0 20.4 ± 1.5 18.7 ± 2.6 • 18.6 ± 0.6 21.5 ± 0.1 21.2 ± 0.8 0.029 .

*: N: P>O.OS; •: O.Ol <P<O.O.S

117

3.2.4 Hypotheses

A number of hypotheses have arisen from the way the Aboriginal communities selected

and sampled tennite mounds. The depth considered in this chapter is depth=! (unless

indicated), which represents the 0-10 em fraction of the outside of the mound. Depth= I

is the depth normally used by Aboriginals. The detailed analysis results of all the

samples are given in Appendices I to V.

3.2.4.1 Hypothesis 1: The New Material of Nasutitermes triodiae Mounds Contains a Higher Element Content, in Particular Iron and Calcium, and Has a Higher Clay and Silt Content than the Older Part of the Mounds.

Samples from new and old material were collected from 5 Nasutitermes triodiae mounds

(n=30), at Daly River (site 4). The results (mean ± standard deviation), per age group

of the mineral analyses (mg/1 OOg) and the particle size analyses (percent of the fine

fraction: <2mm) together with the probability of differences between old and new parts

of the mounds are given in Table 3.11 . The sample depth studied is depth=O, it

represents the 0-1 em fraction collected from the outside of the mound.

The results of the ANOV A show that the only significant differences between the ages

of samples were for aluminium, copper and iron, where the increase in the old material

was highly significant (P<O.OI) and for potassium significantly different (P<O.OS)

(Figure 3.8).

118

5000 ..,

4ooo I a, 0 ;2 3000 i ' � 5 � 2000 -i ()

10] FIGURE 3.8

5000 "")

4000 i a, 0 � 3000 1 ' � 5 S? 2000 -l 0 ()

100: 1 FIGURE 3.9

1.0 12 New marerial

n 1 0 Old rna !erial 1

DB

I

I

I � 0.6

� 04

I I B 02

[)! 0.0 AI ,, K c"

MINERAL MINERAL

Age effects (mean ± SE) on aluminium, iron, potassium and copper (mg/lOOg) in Nasutitermes triodiae mounds sampled in Daly River (site 4)

e.! New malenal Aluminium -

2000 1 Iron 0 Old material

� 1 m

l r,a

I I Top

� _L 1600

l r,a I I,!, 1200

l r,a I P1 BOO

I I I I 400

0 Middle Bottom Top Middle Bottom

POSITION POSITION

Age effects (mean ± SE) on aluminium and iron (mgllOOg), at three positions (top, middle, bottom), in Nasutitermes triodiae mounds sampled in Daly River (site 4)

119

TABLE 3.11 Age effects on selected elements (mgllOOg) and particle sizes (%) (mean ± standard deviation) in 5 Nasutitermes triodiae mounds (Daly River, site 4). ANOVA probability of differences (P) between ages. (Depth=O).

Element I Nasutitermes triodiae site 4 Particle size Old material New material p

n=IS n=JS � Aluminium 3919 ± 6 1 1 3216 ± 380 0.001 ••

Calcium 26.9 ± 10.3 29.2 ± 9.2 0.516 N

Cobalt 0.27 ± 0.06 0.24 ± 0.03 0.092 N

Copper 0.78 ± 0.12 0.67 ± O.o7 0.001 ..

Iron 1364 ± 345 1057 ± ISO 0.004 ••

Potassium 602 ± 35 563 ± 48 0.016 •

Magnesium 95 ± 24 89 ± 10 0.430 N

Manganese 2.12 ± 0.67 2.60 ± 0.70 0.063 N

Sodium 22.7 ± 4.2 21.3 ± 3.0 0.312 N

Zinc 0.46 ± 0.06 0.42 ± 0.06 0.095 N

Clay 19.9 ± 2.9 18.9 ± 3.2 0.382 N

Silt 5.3 ± J.J 5.9 ± 0.6 0.068 N

Fine sand 31.2 ± 2.9 32.9 ± 3.4 0.133 N

Coarse sand 42.0 ± 2.9 40.4 ± 2.3 0.109 N

t: N: P>0.05; •: O.Ol<P<O.OS; .. : O.OOJ<P<O.OI

The aluminium, copper, iron, and to a lesser extent potassium, mean content is

consistently higher in the old part of the mound, at each mound position (top, middle

and bottom), as shown in Figure 3.9 for aluminium and iron. The age effect

probabilities per position in the moWld (Table 3.12) were not significant (P>0.05) in all

cases and the age/position pairwise comparisons (new-old material I top-middle-bottom

section) indicated no significant differences for any of the elements or particle sizes, in

the Nasutitermes triodiae mounds (site 4), except aluminium (Table 3.13). For

aluminium, the pairs top-old versus bottom-new and middle-old versus bottom-new were

significantly different (P<0.05).

TABLE 3.12

Element/ Particle size

Aluminium

Calcium

Cobalt

Copper

Iron

Potassium

Magnesium

Manganese

Sodium

Zinc

Clay

Silt

Fine sand

Coarse sand

*= N: P>0.05

Age (new, old) and position (top, middle, bottom) effects on selected elements (mg/lOOg) and particle size (%) (mean ± standard deviation) in 5 Nasutitermes triodiae mounds sampled at site 4. AN OVA probability of differences (P) between ages. (Depth=O).

Top Middle

Old material New material p Old material New material n=5 n=5 t n=5 n=S

4044 ± 821 3400 ± 325 0.384 N 3995 ± 550 3301 ± 265

22.7 ± 6.6 31.7 ± 9.0 0.677 N 2H ± 7.6 31.5 ± 1 1 .5

0.29 ± 0.08 0.25 ± 0.04 0.892 N 0.27 ± O.o7 0.25 ± O.o3

0.79 ± 0.14 0.71 ± 0.04 0.652 N 0. 79 ± 0.09 0.67 ± 0.05

1411 ± 499 1128 ± 177 0.602 N 1406 ± 352 1080 ± 167

592 ± 53 570 ± 54 0.957 N 6 1 1 ± 23 585 ± 37

92.7 ± 25.0 93.8 ± 7.4 1 .000 N 90.4 ± 20.1 95.3 ± 9.5

1.88 ± 0.39 2.72 ± 0.72 0.410 N 2.00 ± 0.54 2.81 ± 0.77

22.1 ± 5.1 21.6 ± 3.0 1 .000 N 22.1 ± 3.7 22.5 ± 4 . 1

0.46 ± 0.06 0.43 ± O.o3 0.957 N 0.47 ± 0.05 0.43 ± 0.05

19.7 ± 4.1 18.8 ± 1.7 0.998 N 20.4 ± 3.0 18.8 ± 3.6

5.3 ± 0.9 6.4 ± 0.4 0.288 N 5.9 ± 1.4 5.5 ± 0.5

33.8 ± 3.2 33.7 ± 2.2 1 .000 N 30.0 ± 1.2 32.4 ± 4.7

39.7 ± 2.7 39.9 ± 3.4 1 .000 N 43.1 ± 3.0 40.7 ± 2.4

Bottom

p Old material New material p t n=5 n=5 t

0.307 N 3718 ± 505 2947 ± 432 0.208 N

0.894 N 32.9 ± 14.2 24.5 ± 6.3 0.737 N

0.989 N 0.26 ± 0.04 0.22 ± O.o2 0.819 N

0.344 N 0.76 ± 0.09 0.63 ± 0.09 0.192 N

0.452 N 1274 ± 173 964 ± 40 0.508 N

0.916 N 604 ± 28 535 ± 46 0.135 N

0.998 N 102 ± 30 80.0 ± 7.0 0.444 N

0.439 N 2.47 ± 0.94 2.27 ± 0.62 0.997 N

1 .000 N 23.8 ± 3.7 19.8 ± 1.5 0.590 N

0.852 N 0.43 ± 0.06 0.40 ± 0.09 0.946 N

0.970 N 19.7 ± 1.8 19.2 ± 4.5 1 .000 N

0.978 N 4.8 ± 0.7 5.9 ± 0.5 0.344 N

0.808 N 29.6 ± 2.1 32.7 ± 3.5 0.598 N

0.679 N 43.2 ± 2.0 40.5 ± 1.2 0.593 N

� ... =

TABLE 3.13

Position-Age

T·O

T·n

M·o

M·n

B-o

B·n

121

Matrix of Pairwise Comparison Probabilities (P:) (Tukey test) between different positions of material (T=top, M=middle, B=bottom) and age (o=old, n=new) for aluminium content in Nasutitermes triodiae mounds sampled at site 4.

T·o

N

N

N

N

N

•

T·n

N

N

N

N

N

M·o

N

N

N

•

M·n

N

N

N

B·o

N

N

B·n

N :: N: P>0.05; *: O.Ol<P<0.05

3.2.4.2 Hypothesis 2: There is No Difference Between Samples Taken

from Different Positions of Termitaria for

Selected Elements and Particle Size Content.

Three species of termitaria at different sites were analysed separately for position effects:

a) Amitermes vitiosus rnotmds: top and bottom positions;

b) Tumulitermes pastinator mounds: top and bottom positions;

c) Nasutilermes triodiae mounds: top, middle and bottom positions.

A) Amitermes vitiosus

The results of selected elements, particle sizes and ANOV A analyses on the position

effects in Amitermes vitiosus mounds from Daly River (sites 2 and 4) and Elliott (site

5) are given in Table 3.14.

For the Daly River site 2, there were no significant differences (P>0.05) for the ten

selected elements and for particle size composition between the top and the bottom

sections of the mounds. This contrasts with the mounds at Daly River site 4, where the

top section showed a highly significant increase in calcium, copper, magnesium,

TABLE 3.14

Element/

Particle size

Aluminium

Calcium

· Cobalt

Copper

Iron

Potassium

Magnesium

Manganese

Sodium

Zinc

Clay

Silt

Fine sand

Coarse sand

Position effects on selected elements (mgllOOg) and particle size (%)(mean ± standard deviation) in Amitermes vitiosus mounds sampled at sites 2, 4 and 5. ANOV A probability of differences (P) between positions.

Da1y River (Site 2} DaJy River (Site 4) Elliott (Site 5)

Top Bottom Probability Top Bottom Probability Top Bottom Probability n=4 n= 4 t n = lO n= 10 t n=ll n= 1 1 t

3042 ± 143 3209 ± 300 0.353 N 3903 ± 353 3750 ± !59 0.228 N 3771 ± 394 3526 ± 359 0.136 N

39.0 ± [ [ .[ 29.4 ± 7.2 0.!95 N 82.7 ± 27.8 31.8 ± 10.8 0.000 *** [ [7 ± 17 95 ± 20 0.013 .

0.57 ± 0.10 0.55 ± 0.13 0.!61 N 0.50 ± 0.23 0.34 ± O.Dl 0.047 * 0.32 ± 0.05 0.30 ± 0.05 0.372 N

0.83 ± 0.12 0.80 ± 0.17 0.384 N 0.95 ± O.ll 0.79 ± 0.09 0.002 •• 0.93 ± 0.08 0.89 ± 0.07 0.165 N

1574 ± 176 1549 ± 227 0.683 N 1480 ± 341 1359 ± 248 0.376 N 1769 ± 161 [700 ± 141 0.296 N

446 ± 35 469 ± 54 0.874 N 734 ± 82 694 ± 25 0.159 N 165 ± 12 159 ± 12 0.209 N

lOS ± 13 ll3 ± [8 0.776 N 131 ± 17 96 ± 13 0.000 ..... 96 ± 7.1 89 ± 5.9 0.013 .

8.[8 ± 2.36 6.49 ± 1.60 0.!66 N 7.23 ± 1.81 3.88 ± 0.77 0.000 ••• 7.75 ± 1.99 7.21 ± 1.89 0.512 N

12.0 ± 0.5 11.7 ± 2.0 0.998 N 32.6 ± 4.7 28.9 ± 3.4 0.060 N 6.62 ± 0.79 6.35 ± 0.63 0.386 N

0.93 ± 0.16 0.85 ± 0.08 0.277 N 0.55 ± 0.14 0.39 ± 0.05 0.006 • • [.[2 ± 0.08 1.04 ± 0.08 0.026 .

14.7 ± 2.0 16.0 ± 1.6 0.620 N 13.2 ± 4.1 [9.[ ± 3.3 0.002 •• 16.0 ± 6.0 16.7 ± 5.0 0.771 N

14.7 ± 2.2 15.4 ± 2.9 0.145 N 21.4 ± 6.[ 12.0 ± 3.1 0.000 ••• 13.9 ± 2.9 12.9 ± 4.9 0.559 N

28.7 ± 4.7 33.7 ± 3.2 0.899 N 45.1 ± 5.2 41.2 ± 3.4 0.059 N 36.2 ± 4.6 34.9 ± 2.[ 0.410 N

43.8 ± 3.6 36.8 ± 2.8 0.431 N 19.9 ± 5.9 28.6 ± 3.4 0.001 •• 37.2 ± 2.1 38.2 ± 2.0 0.261 N

:1:: N: P>0.05; *: 0.01 <P<0.05; .. : 0.001 <P<O.Ol; ***: P<O.OOI n: number or mounds

� ... ...

123

manganese, zinc and silt; a significant increase in cobalt and a highly significant

decrease in clay and coarse sand, as seen in Figures 3.10 and 3 . 1 1 .

In Elliott, there were significant (P<O.Ol) differences between position for calcium,

magnesium and zinc (Table 3.14). Here, as at Daly River site 4, the increase was in the

top position, with the calcium concentration: 1 1 7 ± 17 mg/1 OOg in the top section and

95 ± 20 mg/IOOg in the bottom section.

In general, there were no highly significant differences between the top and bottom

positions in Amitermes vitiosus mounds for aluminium, cobalt, iron, potassium, sodium

and fine sand content for the three sites studied. For the other elements, the differences

were highly significant but only at site 4.

B) Tumulitermes pastinator

The results of the analyses comparing top with bottom sections of Tumulitermes

pastinator mounds from sites 1 and 6 are given in Table 3.15. The results of the

ANOV A show that the effects of mound position (top/bottom) on the selected elements

and particle size content, were not highly significant (P<0.01) for all the termitaria

studied at the 2 locations. The only significant differences (0.01 <P<O.OS) were for

aluminium, calcium and manganese content at site 1 (Table 3.15).

124

150 ] 10

120 � 8

" 0 0 90 j I F==llil 6 � ' 0 .§ 0 60 j ""' = 4 z 0 0 3: 1 I I 2

0 c. Mg Mo

MINERAL MINERAL

FIGURE 3.10 Position effects (mean ± SE) on calcium, magnesium and manganese (mg/IOOg) in Amitennes vitiosus mounds sampled in Daly River (site 4)

50

40

� 30 w 0 a: 1i: 20

10

O I E Clay Slit Fine sand

PARTICLE SIZE

� Bottom � Top

Coarse sand

FIGURE 3.11 Position effects (mean ± SE) on particle size ( (%) in Amitermes vitiosus mounds sampled in Daly River (site 4)

125

TABLE 3.15 Position effects on selected elements (mgllOOg) and particle sizes (%) (mean ± standard deviation) in Tumulitermes pastinator mounds sampled at sites 1 and 6. ANOV A probability of differences (P) between positions.

Element/ Daly River (Site I)

Particle size Top Bottom n=l5 n=Il

Aluminium 3 1 19 ± 176 2942 ± 196

Calcium 21.4 ± 5.2 26.7 ± 7.8

Cobalt 0.29 ± 0.01 0.29 ± 0.01

Copper 0.57 ± 0.03 0.57 ± O.o3

Iron 1359 ± 101 1321 ± 77

Potassium 514 ± 23 503 ± 19

Magnesium 64.3 ± 7.4 65.7 ± 6.6

Manganese 3.16 ± 0.32 3.57 ± 0.46

Sodium 27.8 ± 2.5 26.4 ± 1.6

Zinc 0.48 ± 0.03 0.48 ± 0.03

Clay 14.8 ± 1.3 14.8 ± 1.4

Silt 8.7 ± 1.0 8.9 ± 1.3

Fine sand 43.3 ± 3.1 43.8 ± 2.0

Coarse sand 32.7 ± 3.7 32.8 ± 2.6

l: N: P>0.05; •: O.Ol<P<O.OS n: number of mounds

C) Nasutitermes triodiae

Howard Springs (Site 6)

p Top Bottom p I n=I3 n=9 I

0.024 • 6300 ± 703 6125 ± 524 0.532 N

0.048 . 52 ± 12 47 ± 10 0.373 N

0.376 N 0.92 ± 0.23 0.84 ± 0.26 0.463 N

0.691 N 1.67 ± 0.33 1.65 ± 0.39 0.881 N

0.302 N 4307 ± 658 4813 ± 971 0.160 N

0.185 N 38.1 ± 7.7 35.6 ± 6.4 0.449 N

0.628 N 48.9 ± 5.3 48.9 ± 4.8 0.975 N

0.014 "' 6.54 ± 1.89 6.61 ± 2.04 0.938 N

0.116 N 6.11 ± 0.66 6.45 ± 0.67 0.258 N

0.997 N 1.14 ± 0.16 1.16 ± 0.25 0.876 N

0.999 N 27.3 .± 3.4 25.8 ± 3.5 0.322 N

0.560 N 6.41 ± 1.0 5.9 ± 1.1 0.358 N

0.605 N 45.9 ± 4.3 46.7 ± 2.2 0.647 N

0.967 N 18.8 ± 2.3 19.8 ± 3.3 0.405 N

The results of the analyses comparing the top and bottom positions of Nasutitermes

triodiae mounds, at sites 4 and 6 are given in Table 3.16. As for Tumulitermes

pastinator, the results of the AN OVA show no highly significant differences between

mound positions for any of the elements or particle sizes. The only significant

differences (O.Ol<P<0.05) were in the site 4 samples, with a significant increase in

aluminium and copper in the top section of the mounds. No position effects were

observed in the Howard Springs mounds.

� TABLE 3.16 Position effects on selected elements (mg/lOOg) and particle size (o/o) (mean ± standard deviation) in Nasutitermes

... a-triodiae mounds sampled at sites 4 and 6. ANOV A probability of differences (P) between positions.

Element/ Daly River (Site 4) Howard Springs (site 6)

Particle size Top Middle Bottom Probability Top Middle Bottom Probability n=l5 n=IO n= l5 * n=S n=S n=S *

Aluminium 4150 ± 539 4327 ± 694 3735 ± 407 0.020 • 5774 ± 761 5723 ± 892 5716 ± 708 0.992 N

Calcium 41.4 ± 22.6 45.5 ± 26.4 38.3 ± 23.3 0.755 N 85 ± 19 82 ± 27 84 ± 28 0.981 N '

Cobalt 0.34 ± 0.04 0.35 ± 0.05 0.32 ± O.DJ 0.162 N 0.81 ± 0.25 0.78 ± 0.23 0.78 ± 0.21 0.972 N

Copper 0.85 ± 0.09 0.87 ± 0.10 0.78 ± 0.07 0.025 • 1.61 ± 0.22 1.54 ± 0.15 1.53 ± 0.15 0.729 N

Iron 1504 ± 333 1600 ± 324 1346 ± 227 0.094 N 3683 ± 406 3624 ± 770 3639 ± 1120 0.993 N

Potassium 668 ± 68 695 ± 85 632 ± 45 0.059 N 38.0 ± 3.7 38.3 ± 5.5 39.2 ± 5.4 0.926 N

Magnesium 103 ± 13 1 15 ± 21 102 ± 22 0.232 N 50.2 ± 6.7 48.8 ± 8.6 52.0 ± 8.9 0.821 N

Manganese 3.63 ± 1.17 3.99 ± 1 .29 3.52 ± 0.68 0.529 N 4.88 ± 1.38 5.08 ± 1.83 5.14 ± 2.14 0.972 N

Sodium 25.5 ± 3.1 28.6 ± 4.5 26.4 ± 3.7 0.126 N 5.73 ± 0.61 5.65 ± 0.68 6.30 ± 1.03 0.394 N

Zinc 0.47 ± 0.08 0.46 ± 0.07 0.42 ± 0.05 0.127 N 1.12 ± 0.13 1.01 ± 0.21 0.98 ± 0.12 0.360 N

Clay 22.1 ± 2.8 23.8 ± 3.9 20.8 ± 2.4 0.054 N 26.8 ± 2.5 28.4 ± 3.2 27.7 ± 3.8 0.732 N

Silt 7.8 ± 2.4 8.9 ± 4.2 7.4 ± 1.8 0.433 N 8.3 ± 1.7 7.8 ± 1.2 7.5 ± 0.7 0.629 N

Fine sand 31.7 ± 3.2 32.0 ± 4.7 34.7 ± 3.5 0.071 N 43.9 ± 3.0 44.7 ± 1.9 45.5 ± 4.2 0.730 N

Coarse sand 38.6 ± 3.4 35.1 ± 6.4 36.3 ± 4.2 0.168 N 20.8 ± 2.6 18.2 ± 3.0 18.6 ± 1.2 0.221 N

�: N: P>0.05; •: 0.01 <P<O.OS n: number of mounds

3.2.4.3 Hypothesis 3:

127

There are No Significant Selected Elements and

Particle Size Differences Between Mounds of

Different Sizes.

To test the influence of the mound size on selected elements and particle sizes, 10 to 15

Amitermes vitiosus, Tumulitermes pastinator and Nasutitermes triodiae mounds were

sampled at two locations per species. The correlation between the size of the mounds

and the selected elements and particle sizes was calculated by the Pearson Correlation

and Probability Test. Equality of variance and normality were checked by Bartlett's

Test. The results of the analyses, given in Table 3.17, indicate that 85 % of the variable

correlations are not significantly correlated. The significant 15 % of correlations are not

consistent across species and sites. For example, calcium is positively correlated to size

only in Amitermes vitiosus mounds at site 5; iron is not significantly correlated for the

three species at all sites and clay is significantly positively correlated in Nasutitermes

triodiae mounds at site 4. There is no consistency in. the way the variables are

correlated. For example in Amitermes vitiosus mounds the iron is positively correlated

(but not significantly) to size at site 4, while it is significantly negatively correlated to

size at site 5. Overall, there are no significant correlations between mound size and

selected element and particle sizes.

3.2.4.4 Hypothesis 4: There are Differences Between Mounds of the

Same Species at the Same Site.

The differences between mounds of the same species at the same site were tested by

ANOV A for Amitermes vitiosus at sites 2, 4 and 5, Tumulitermes pastinator at sites 1

and 6, Nasutitermes triodiae at sites 4, 6 and 7 and for Tumulitermes hastilis at site I .

The probability differences (P) are given in Table 3.18. The concentration data are

given in Appendices I to V. Interestingly, at two sites (Amitermes vitiosus site 4 and

Tumulitermes pastinator site 1), there were no significant differences between mounds

of the same species, except for iron at site 4 (0.01 <P<0.05), but significant and highly

significant differences for the other species on the same site (Nasutitermes triodiae at site

4 and Tumulitermes hastilis at site 1). An example of the variation between mounds is

128

120 2500

100

� 80 � ' 0 .§ 60

........ � 2000 0 0 0 � 1500 .§

0 15 0 40

� 1000 0

• 0 20

l � § s § § .1: 500

0 o ' ''''''''''''''' 1 � 3 • e o r a a w •• n ffl u � • 2 3 • a e r e e � ,, 12 13 u �

Nt MOUND SITE 4 Nt MOUND SITE 4

FIGURE 3.12 Mound effects (mean ± SE) on calcium and iron (mg/lOOg) in Nasutitermes triodiae

mounds sampled in Daly River (site 4)

129

given in Figure 3.12, for calcium and iron content in Nasutilermes triodiae mounds at

site 4. At the other sites, the differences between mounds are often highly significant

for all the selected elements.

For the particle sizes, the differences are less marked. There was only one site with

highly significant differences for silt.

TABLE 3.17 Pearson correlation (PC) and probability (P�) matrix of mound size (height + circumference) with selected elements and particle sizes of three species® at different sites.

Element/ Amitermes vitiosus Tumulitermes pastinator Nasutitermes triodiae Particle size "s"'it:::e-;4:---;S;:i::te�5;;-----;:S'=ite::-;l---;S;:it::e-;6,---�S;:i::te:-4:;----;;S'=it:::e-.:6--

Size P Size P Size P Size P Size P Size P

Size 1.000 • • • 1.000 • • • 1.000 ••• 1 .000 • • • 1.000 ••• 1 .000 • • •

0.559 *** 0.106 N Aluminium 0.249 N -0.359 N -0.046 N 0.179 N

Calcium

Cobalt

Copper

Iron

0.389 N

-0.068 N

0.341 N

0.320 N

Potassium 0.454 •

Magnesium 0.292 N

Manganese 0.421 N

Sodium 0.295 N

0.527 • •

0.154 N

0.055 N

-0.367 N

-0.3 1 8 N

0.131 N

0.146 N

-0.282 N

-0.087 N

-0.645 • • •

-0.289 N

0.224 N

-0.093 N

-0. 1 1 5 N

-0.567 • •

-0.034 N

0.149 N

0.207 N

0.313 •

0.202 N

0.132 N

0.3 1 5 N

-0.030 N

0.236 N

0.404 ** 0.092 N

0.306 N 0.165 N

0.161 N 0.192 N

0.387 * 0.092 N

Zinc 0.477 • -0.063 N

0.213 N

0.020 N

0.150 N

0.013 N

0,078 N

0.376 N

0.1 18 N

-0.094 N

0.192 N

0.547 N

-0.279 N

-0.164 N 0.294 N -0.282 N Clay -0.019 N

Silt 0.233 N

F.sand 0.031 N

Coarse sand -0.175 N

0.149 N

-0.443 •

-0.016 N

O.oJ8 N

-0.527 N

0.428 N

0.200 N

-0.154 N

-0.610 •••

0.426 •

0.321 * 0.0 1 1 N

0.086 N -0.195 N

-0.045 N -0.199 N

-0.239 N 0.101 N :t N: P>0.05; *: O.OI<P<O.OS; n: O.OOI<P<O.OI; •n: P<O.OOI

@ Amitermes viti=: Tumulitermes pastinator: Nasutllerme.s triodiae:

site 4: 10 mounds (n=20) site 5: I I mounds (n=26) site I: IS mounds (n=26) site 6: 13 mounds (11'"'25)

site 4: IS mounds (n=41) site 6: 5 mounds (n=J5)

130

TABLE 3.18 Probability differences (ANOVA) PW between mounds of each species (Av, Tp, Nt and Th) per site for selected elements and particle sizes.

Element/ Av Tp Nt Th -

Particle size Site 2 Site 4 Site 5 Site I Site 6 Site 4 Site 6 Site 7 Site I n=12 n=20 n=26 n=26 n=25 n=41 n=15 n::JO n:IO

Aluminium • N .. N .. .. .. . .. ...

Calcium N N .. N ... ... ... ... ...

Cobalt ... N .. N ... .. ... .. ..

Copper ... N .. N ... .. .. • ...

Iron .. • ... N • ... .. ... N

Potassium .. N .. N .. N .. N N

Magnesium • N N N ... .. ... .. ...

Manganese • N ... N ... ... ... ... ...

Sodium • N .. N ... • N • N

Zinc N N N N .. ... .. N ...

Clay . . N N N • .. ...

Silt .. N N N N N N

Fine sand N N N N .. .. N

Coarse sand N N ... N .. • N £ N: P>0.05; 1: O.OI<P<O.OS; ": O.OOI<P<O.Ol; n•: P<O.OOI

3.2.4.5 Hypothesis 5: There are Differences in Selected Elements and Particle Sizes Between Termitaria of Different Species at the Same Site.

The data were analysed statistically by one-way analysis of variance (ANOV A), with

selected elements and particle size distribution as the source of variance between species

mounds. An overview of the termitaria selected elements and particle sizes (mean ±

standard error), per species and per site, is presented in Figures 3.13 (A to D).

7000

6000

- 5000 8 2 4000

g 3000 ;;: 2000

1000

0

150

"' 100 8 ' 0 .§ • 50 0

Av

2 4 '

Av

2 4 '

131

Nt Th

3 6 3 4 6 7 SITE

!'!! Th

7

TABLE 3.13 A Selected elements (mg!IOOg) and particle size (%) (mean ± SE) of Amitennes vitiosus

(A v), Tumulitermes pastinator (Tp ), Nasutitermes triodiae (Nt) and Tumulitermes hastilis

(Th) at sites 1-7.

Aluminium and calcium concentrations (mgllOOg)

1.00

OEO 0 8 060 � ' rn ..s 0.40 0 0

0.20

000

2.00 l 150

0 0 0 � � 1.00 s , 0

0.50

000 I

AY In I'!! Th 5000 J Av I In ., , Nt 4000

0 8 3000 � ' � s 2000 m "-

1000

0

2 4 5 I 3 6 3 4 6 7 I 2 4 5 I 3 6 3 4 6

1000 Av I In � I Nt I Th l Av I In ' ""' Nt

1 ..+, BOO

0 0 600

- � 0 �

� � - � I "' s 400

rlil "'

200

vp 9 r�(J '''(' ¥(4 r\Q II"(I !Cf' Kie "I" ' I 0 2 4 5 I 3 6 3 4 6 7 I I 3 6 3 4 6 2 4 5

SITE SITE

TABLE 3.13 B Selected elements (mg/IOOg) and particle size(%) (mean ± SE) of Amilermes vitiosus

(A v), Tumulitermes pastinaror(Tp), Nasutitermes triodiae (Nt) and Tumulitermes hastilis

(Th) at sites 1�7. Cobalt, copper, iron and potassium concentrations (mg/IOOg)

hh I � "' N

7

I Th

7

300 .., I I� I Th I 40 Av In Nt

30 roo ] I -a 8

I -

E3 r 20 � 100 El El • = z 10 J i I iii i �li i � ;li I 0

2 4 5 I 3 6 3 4 6 . 7 I 2 4 5 I 3 6 3 4 6 7

10 , 2.00 Av In Rl Nt Th

8 1.50 -a -a 0 6 I

0 0

I 0 -

� - � -' '(» 1.00 0 s 4 5

c

� c " N 2 0.50

0 ')" "I'' "�' I "j � ·y· ,,, ,,�. "j¥ , ... .,,,. . "i' 0.00 2 4 5 I 3 6 3 4 6 7 I 2 4 5 I 3 6 3 4 6 7

SITE SITE

TABLE 3.13 C Selected elements (mg/IOOg) and particle size(%) (mean ± SE) of Amitermes vitiosus (A v), Tumulitermes pastinator(Tp), Nasutitermes triodiae (Nt) and Tumulitermes hastilis

(Th) at sites 1-7. � "' "' Magnesium, manganese, sodium and zinc concentrations (mg/100g)

so , � 20 l 10

J so , : 20 l 10

0 '

Av I rn ""'I

I I � 1 l l ; Nt J"'l I Th I

� i i iii i �� I

2 4 ' I 3 6 3 4 6 7 '

Av I In I� Nt I Th I

� - � = 1 � � � r?l � Yf' Yf" •1a "'{' Kf' 'j" "'J" "(' "I" ' '

2 4 ' I 3 6 3 4 6 7 I

SJT�

60 l I� rn �1 Av ---40

g 30 " � 20 u_

10

0 2 4 ' I 3 6 3

60 l Av I In I

40

g so " • c.Q 20 "

10

0 2 4 ' I 3 6 3

SITE

TABLE 3.13 D Selected elements (mg/IOOg) and particle size(%) (mean ± SE) of Amitennes vitiosus

Nt � � I !" I

4 6 7

Nt

4 ' 7

(A v), Tumulitermes pastinator(Tp), Nasurilermes triodiae (Nt) and Tumulitermes hastilis

(Th) at sites 1-7. Clay, silt, fine sand and coarse sand content (%)

-... ...

135

A) Tumulitermes pastinator versus Tumulitermes hastilis mounds at site 1 (Daly

River).

The mean (± standard deviation) of the selected elements, the particle size distribution

and the pairwise comparison probabilities between Tumulitermes pastinator and

Tumulitermes hasti/is mounds, sampled at site 1 , are given in Table 3.19.

As seen in Table 3.19, the aluminium, potassium, sodium, zinc, clay and to a lesser

extent fine sand content are highly significantly increased (P<O.Ol) in Tumulitermes

pastinator mounds, while the calcium, magnesium, manganese and coarse sand content

are higher in Tumulitermes hastilis mounds. For example, in Tumulitermes pastinator

motu1ds: aluminium is 38 % higher; calcium 4.5 fold lower and coarse sand 22 % lower.

For the other selected elements and particle sizes (cobalt, copper, iron and fine sand)

there were no significant differences detected at site 1 .

136

TABLE 3.19 Selected elements (mg/100g) and particle sizes (%) (mean ± standard deviation) of Tumulitermes pastinator and Tumulitermes hastilis mounds sampled at site 1. Probability of differences (P) between the two species mounds@.

Element/ Tumulitermes Tumulitermes p Particle size pas tina tor hastilis �

n=26 n=S

Aluminium 3044 ± 201 2209 ± 247 • •••

Calcium 23.7 ± 6.8 107 ± 66 ...

Cobalt 0.29 ± 0.01 0.30 ± 0.04 N

Copper 0.57 ± 0.03 0.57 ± 0.07 N

Iron 1343 ± 92 1249 ± 1 13 • N

Potassium 510 ± 22 403 ± 18 • ...

Magnesium 64.9 ± 6.9 76 ± 14.2 ..

Manganese 3.33 ± 0.43 5.91 ± 1 .65 ...

Sodium 27.2 ± 2.2 18 .4 ± 1 .02 • ...

Zinc 0.48 ± 0.03 0.42 ± 0.07 • ..

Clay 14.8 ± 1.3 12.3 ± 2.6 • ..

Silt 8.8 ± 1.1 9.6 ± 0.6 N

Fine sand 43.5 ± 2.7 38.9 ± 10.1 • •

Coarse sand 32.8 ± 3.2 42.0 ± 1 1 . 1 ...

@: 15 Tumulitermes pastinator mourids and Tumulilermes hastilis mounds

I ' N: P>O.OS; $; O.Ol<P<O.OS; U; O.OOI<P<O.Ol; •••: P<O.OOI .o. : mean variable content higher in Tunrulitermu pastinator mounds

B) Tumu/itermes pastinator versus Nasutitermes triodiae mounds at site 3 (Daly

River).



Nasutitermes triodiae mounds, sampled at site 1 , are given in Table 3.20.

The probability of differences (ANOV A) between the two species' mounds indicates a

significant increase of fine sand (P<O.OS) and a highly significant increase of coarse sand

(P<O.OOl) in Tumulitermes pastinator mounds. However, although the copper content

is 1.50 ± 0.55 mg/IOOg in Tumulitermes pastinator mounds and 0.97 ± 0.30 mgl100g

137

m Nasutitermes triodiae mounds, no significant difference was indicated. In

Nasutitermes triodiae, there is a highly significant increase in aluminium, cobalt, iron,

potassium, magnesium, manganese and silt content (P<O.Ol) and a significant increase

of sodium (P<O.OS). For example, the mean magnesium and manganese content are

respectively 69 and 57 % higher in Nasutitermes triodiae.

There are no significant statistical differences detected at site 3 for calcium, copper, zinc

and clay content between the two species.

TABLE 3.20 Selected elements (mg/IOOg) and particle sizes (%) (mean ± standard deviation) of Tumulitermes pastinator and Nasutitermes triodiae mounds sampled at site 3. Probability of differences (P) between the two species mounds®.

Element/ Tumulitermes Nasutitermes p Particle size pastinator triodiae t

n=6 n=6

Aluminium 3826 ± 446 4488 ± 167 ••

Calcium 23.3 ± 3.9 23.1 ± 4.72 • N

Cobalt 0.42 ± 0.04 0.52 ± 0.04 ••

Copper 1.50 ± 0.55 0.97 ± 0.30 • N

Iron 2917 ± 198 3434 ± 124 ...

Potassium 651 ± 79 897 ± 65 •••

Magnesium 1 5 1 ± 19 256 ± 24.9 •••

Manganese 5.97 ± 0.83 9.38 ± 0.99 •••

Sodium 13.9 ± 2.1 16.5 ± 2.03 •

Zinc 1.54 ± 0.28 1.68 ± 0.18 N

Clay 21.2 ± 2.4 19.4 ± 1.0 • N

Silt 15.6 ± 2.2 29.4 ± 1.3 ...

Fine sand 36.9 ± 2.1 33.6 ± 2.1 • •

Coarse sand 28.8 ± 2.9 20.4 ± 1.5 • ...

@ : I Tumuliterm£s pastinator mound and I Nasutitermes triodiae mound

I' N: P>O.OS; *: O.Ol<P<O.OS; .. : O.OOI<P<O.Oi; •••: P<O.OOI " mean variable content higher in Tumulitermes pastinator mounds

138

TABLE 3.21 Selected elements (mg/100g) and particle sizes (%) (mean ± standard

deviation) of Amitermes vitiosus and Nasutitermes triodiae mounds

sampled at site 4. Probability of differences (P) between species

mounds®.

Element/ Amitermes Nasutitermes p Particle size vitiosus triodiae �

n�20 n=ll

Aluminium 3826 ± 278 4031 ± 578 N Calcium 57.2 ± 33.2 41.2 ± 23.4 • •

Cobalt 0.42 ± 0.18 0.33 ± 0.04 • ..

Copper 0.87 ± 0.13 0.83 ± 0.09 • N Iron 1420 ± 297 1466 ± 304 N Potassium 714 ± 62 660 ± 68 • ..

Magnesium 1 14 ± 23 106 ± 19 • N

Manganese 5.56 ± 2.19 3.67 ± 1.03 • ...

Sodium 30.8 ± 4.4 26.6 ± 3.8 • ...

Zinc 0.47 ± 0.13 0.45 ± 0.07 • N Clay 16.2 ± 4.8 22.03 ± 3.1 ...

Silt _16.7 ± 6.8 7.9 ± 2.8 • ...

Fine sand 43.2 ± 4.7 32.9 ± 2.9 • ...

Coarse sand 24.3 ± 6.4 36.9 ± 4.7 •••

@: 10 A.mitermes vitioswr mounds and IS Nasulitermes triodiae mounds

!: N: P>O.OS; •: O.Oi<P<O.OS; �•: O.OOI<P<O.Oi; •n: P<O.OOI & : mean variable content higher in Amilermes vitiosus

C) Amitermes vitiosus versus Nasutitermes triodiae mounds at site 4.

139


and the probabilities (P) between Amitermes vitiosus and Nasutitermes triadiae mounds,

sampled at site 4, are presented in Table 3.21.

The results of the ANOV A betweenAmitermes vitiosus and Nasutitermes triodiae mound

contents, indicate highly significant differences (P<O.Ol ) for cobalt, potassium,

manganese, sodium, clay, silt, fine sand and coarse sand and a significant di�erence

(?<0.05) in calcium content. Nasutitermes triodiae mounds have a larger proportion of

clay (36 % higher) and coarse sand (52 % higher) than the Amitermes vitiosus mounds.

Amitermes vitiosus mounds have a higher calcium, cobalt, potassium, manganese (51 %),

sodium, silt (1 1 1 %) and fine sand (%) content than Nasutitermes triodiae mounds.

No significant differences were observed for aluminium, copper, iron, magnesium and

zinc content between the Amitermes vitiosus and Nasutitermes triodiae mounds studied

at site 4.

D) Tumulitermes pastinator versus Nasutitermes triodiae mounds at site 6 (Howard Springs).



Nasutitermes triodiae mounds sampled at site 6, are given in Table 3.22.

Significant differences are observed between the two species for: aluminium (P<0.05),

calcium (P<O.OOl), iron (P<O.Ol), manganese (P<0.05) and silt (P<O.OOI). No

significant differences are observed for the other variables.

140

TABLE 3.22 Selected elements (mgllOOg) and particle sizes(%) (mean ± standard

deviation) of Tumulitermes pastinator and Nasutitermes triodiae

mounds� sampled at site 6. Probability of differences (P) between the

3.2.4.6

two species mounds.

Element/ Tumu/itermes Nasutitermes Particle size pas tina/or triodiae

n�25 0""15

Aluminium 6300 ± 658 5738 ± 733

Calcium 50.0 ± I I . 7 83.7 ± 23.2

Cobalt 0.86 ± 0.24 0.79 ± 0.21

Copper 1 .63 ± 0.35 !.56 ± 0.17

Iron 4527 ± 780 3649 ± 759

Potassium 37.3 ± 7.4 38.5 ± 4.6

Magnesium 49.3 ± 5.6 50.3 ± 7.62

Manganese 6.40 ± 1.88 5.03 ± 1.68

Sodium 6.22 ± 0.73 5.89 ± 0.79

Zinc 1 . 15 ± 0.22 1.04 ± 0.16

Clay 26.9 ± 3.5 27.6 ± 3.0

Silt 629 ± 1 .1 7.9 ± 1 2

Fine sand 45.6 ± 3.8 44.7 ± 3.0

Coarse sand 19.5 ± 2.9 19.2 ± 2.5

fl: 13 Tumulitermes pastinator mounds and S Nasuli/ermes triodiae mounds

N: P>0.05; •: O.Ol<P<0.05; n: O.OOI<P<O.OI; u•: P<O.OOI .o : mean variable content higher in Tumulitermes pastinator mounds

•

•

•

•

p t

•

...

N

N ..

N

N

• •

• N

• N

N ...

• N

• N

Hypothesis 6: There are Differences in Element and Particle

Size Content for Same Species Mounds at Different Sites.

A) Amitermes vitiosus

The mean (± standard deviation) of selected elements (mg/IOOg) and particle sizes (%)

of Amitermes vitiosus mounds at sites 2, 4, 5 and at all sites (2+4+5) are presented in

Table 3.23 together with the probability of differences (P) between the Amitermes

vitiosus termitaria samples collected at the three sites.

141

TABLE 3.23 Selected elements (mg/100g) and particle sizes (%) (mean ± standard deviation) of Amitermes vitiosus mounds per site (2, 4 and 5) and for all sites (2+4+5) together with the probability of differences (P) between sites.

Element/ Daly River Daly River Elliott Total p Particle size (Site 2) (Site 4) (Site 5) (Sites 2+4+5) j

n=12 n=20 n=26 n=58

Aluminium 3 1 15 ± 193 3826 ± 278 3634 ± 371 3593 ± 402 ...

Calcium 32.0 ± 9.9 57.2 ± 332 1 1 1 ± 23.9 76.2 ± 41.6 ...

Cobalt 0.55 ± 0.11 0.42 ± 0.18 0.31 ± 0.04 0.40 ± 0.15 ...

Copper 0.80 ± 0.14 0.87 ± 0.13 0.92 ± O.o7 0.88 ± 0.12 •

Iron 1521 ± 198 1420 ± 297 1726 ± 146 1578 ± 256 ...

Potassium 455 ± 38 714 ± 62 161 ± 12 413 ± 251 ...

Magnesium 106 ±. 14 1 14 ± 23 93 ± 7 103 ± 1 8 ...

Manganese 6.92 ± 1.99 5.56 ± 2.19 7.54 ± 1.81 6.73 ± 2.13 ..

Sodium 1 1.7 ± 1.22 30.8 ± 4.4 6.47 ± 0.68 15.9 ± 1 1.4 ...

Zinc 0.89 ± 0.14 0.47 ± 0.13 1.09 ± 0.08 0.83 ± 0.30 ...

Clay 15.4 ± 1.7 16.2 ± 4.8 16.7 ± 5.4 16.3 ± 4.6 N

Silt 14.8 ± 2.3 16.7 ± 6.8 12.9 ± 4.2 14.6 ± 52 •

Fine sand 31.8 ± 4.2 43.2 ± 4.7 352 ± 3.6 37.2 ± 6.1 ...

Coarse sand 40.1 ± 4.5 24.3 ± 6.4 37.8 ± 2.0 33.6 ± 8.2 ...

t: N: P>0.05; •: O.Ol<P<O.OS; n: O.OOI<P<O.OI; •••: P<O.OOI

The ANOVA shows that there are highly significant differences (P<O.OOl) in nearly all

the variables studied: aluminium, calcium, cobalt, iron, potassium, magnesium,

manganese, sodium, zinc, fine sand and coarse sand; significant differences (P<0.05) in

copper and silt and no significant difference (P>O.OS) in clay content (the mean clay

content remained at approximately 16 %). The aluminium, copper, iron, magnesium,

manganese and fine sand mean content, although highly significantly different, varied

within a narrow range. This contrasts with the mean contents of calcium, potassium,

sodium, zinc and coarse sand, which vary greatly between sites (Figure 3.13, A to N ).

142

B) Tumulitermes pastinator

The mean (± standard deviation) of selected elements (mg/1 OOg) and particle sizes

analyses (%) of Tumulitermes pastinator mounds at sites 1 , 3, 6 and at all sites (1+3+6)

are presented in Table 3.24, together with the probability of differences (P) between all

Tumulitermes pastinator samples collected.

The AN OVA shows that there are highly significant differences P(<O.OOl) between sites

for all the selected elements and particle sizes. Figure 3.13 (A to N) shows the wide

variation between sites. For example, the mean content varied from 3044 ± 201 to 6300

± 658 mgllOOg for aluminium, 1343 ± 92 to 4527 ± 780 mg/lOOg for iron and 27.2 ±

2.2 to 6.22 ± 0.73 mgllOOg for sodium.

TABLE 3.24 Selected elements (mg/IOOg) and particle sizes (%) (mean ± standard deviation) of Tumulitermes pastinator mounds per site (1, 3 and 6) and for all sites (1+3+6) together with the probability of differences (P) between sites.

Element/ Daly River Daly River Howard Springs Total Probability Particle (Site I) (Site 3) (Site 6) (Sites 1+3+6) P t SIZe n�26 n=6 n�5 n�57

Aluminium 3044 ± 201 3826 ± 446 6300 ± 658 4554 ± 1643 •••

Calcium 23.7 ± 6.8 23.3 ± 3.9 50.0 ± 1 1 .7 35.2 ± 15.9 ...

Cobalt 0.29 ± O.Ql 0.42 ± 0.04 0.86 ± 0.24 0.55 ± 0.32 ...

Copper 0.57 ± 0.03 1.50 ± 0.55 1.63 ± 0.35 1.13 ± 0.59 ...

Iron 1343 ± 92 2917 ± 198 4527 ± 780 2905 ± 1605 ...

Potassium 510 ± 22 651 ± 79 37.3 ± 7.4 3 1 8 ± 255 ...

Magnesium 64.9 ± 6.9 151 ± 19 49.3 ± 5.6 67.2 ± 3 1 .3 ...

Manganese 3.33 ± 0.43 5.97 ± 0.83 6.40 ± 1.88 4.96 ± 1.98 ...

Sodium 27.2 ± 2.2 13.9 ± 2.1 6.22 ± 0.73 16.6 ± 10.2 ...

Zinc 0.48 ± 0.03 1.54 ± 0.28 1 .15 ± 0.22 0.89 ± 0.42 ...

Clay 14.8 ± 1.3 21.2 ± 2.4 26.9 ± 3.5 20.8 ± 6.2 •••

Silt 8.8 ± 1.1 15.6 ± 2.2 6.29 ± 1.1 8.4 ± 3.0 ...

Fine sand 43.5 ± 2.7 36.9 ± 2.1 45.6 ± 3.8 43.7 ± 4.1 •••

C.sand 32.8 ± 3.2 28.8 ± 2.9 !9.5 ± 2.9 26.5 ± 7.1 *** l: •••: P<O.OOl

143

Most of the lowest mean values are found in the Daly River site I samples (except for

potassium, sodium and coarse sand); while the highest mean values are found in the

Howard Springs samples. Site 3 has a particularly high copper, potassium, magnesium,

manganese, zinc and silt mean content.

C) Nasutitermes triodiae

The mean (± standard deviation) of selected elements (mg/l OOg) and particle sizes (%)

of Nasutitermes triodiae mounds per site (3, 4, 6 and 7) and for all sites (3+4+6+ 7) are

presented in Table 3.25 together with the probability of differences (P) between all the

Nasutitermes triodiae samples collected.

The ANOV A shows that like the Tumulitermes pastinator mean contents, there are

highly significant differences between sample content for all the selected elements and

particle sizes. Figure 3.13 (A to N) shows the wide variation between sites. For

example, the mean content varied from: 23.1 ± 4.7 to 83.7 ± 23.2 mg/100g for calcium,

890 ± 134 to 3649 ± 759 mg/1 OOg for iron and 38.5 ± 4.6 to 897 ± 65 mgllOOg for

potassium. The variation between clay content remained small: 19.2 ± 5.9 % to 27.6

± 3.0 %

144

TABLE 3.25 Selected elements (mg/IOOg) and particle sizes (%) (mean ± standard deviation) of Nasutitermes triodiae mound samples per site (3, 4, 6 and 7) and for all sites (3+4+6+7) together with the probability of differences (P) between sites.

Element/ Daly River Daly River Howard Berrimah Total (Sites P l Particle size (Site 3) (Site 4) Springs (Site 7) 3+4+6+7)

n=6 n-41 (Site 6) n=S n=67 n=15

Aluminium 4488 ± 167 4031 ± 578 5738 ± 733 3330 ± 337 4402 ± 951 ••• Calcium 23.1 ± 4.72 41 .2 ± 23.4 83.7 ± 23.2 82.4 ± 28.2 52.2 ± 30.7 ••• Cobalt 0.52 ± 0.04 0.33 ± 0.04 0.79 ± 0.21 0.34 ± O.QJ 0.45 ± 0.22 ••• Copper 0.97 ± 0.30 0.83 ± 0.09 1.56 ± 0.17 0.89 ± 0.04 1.01 ± 0.33 ••• Iron 3434 ± 124 1466 ± 304 3649 ± 759 890 ± 134 2088 ± 1 1 17 ••• Potassium 897 ± 65 660 ± 68 38.5 ± 4.6 341 ± 19 519 ± 289 ••• Magnesium 256 ± 24.9 106 ± 19 50.3 ± 7.62 58.8 ± 5.8 103 ± 57 ••• Manganese 9.38 ± 0.99 3.67 ± 1.03 5.03 ± 1.68 3.19 ± 0.55 4.45 ± 2.03 ••• Sodium 16.5 ± 2.03 26.6 ± 3.8 5.89 ± 0.79 33.4 ± 3.7 21.6 ± 9.73 ••• Zinc 1.68 ± 0.18 0.45 ± O.Q7 1 .04 ± 0.16 0.64 ± 0.05 0.71 ± 0.41 ••• Clay 19.4 ± 1.0 22.0 ± 3.1 27.6 ± 3.0 19.2 ± 5.9 22.8 ± 4.2 ••• Silt 29.4 ± 1.3 7.9 ± 2.8 7.9 ± 1.2 10.0 ± 2.0 10.0 ± 6.6 ••• Fine sand 33.6 ± 2.1 32.9 ± 2.9 44.7 ± 3.0 45.3 ± 1.8 36.6 ± 6.4 ••• Coarse sand 2o.4 . ± 1.5 36.9 ± 4.7 19.2 ± 2.5 22.3 ± 4.6 30.4 ± 9.2 •••

t *-*: P<O.OOI

The Howard Springs site had the highest aluminium, calcium, cobalt, copper, iron, clay

and fine sand content; while the higher potassium, magnesium, manganese, sodium, zinc,

silt and coarse sand mean values were found principally at the Daly River, site 3. The

iron content is exceptionally low at the Berrimah site while the sodium content is the

highest.

3.2.4. 7 Hypothesis 7:

145

There are Differences in Elements and Particle

Sizes Between Different Species at Different

Sites.

The minimum, maximum and mean (± standard deviation) of selected elements

(mg/l OOg) and particle sizes (%) of the species of mounds (Amitermes vitiosus,

Tumulitermes pastinator and Nasutitennes triodiae) chosen by the Aboriginal

communities, sampled at sites (1 to 7), together with the probability of differences

between the three species, are given in Table 3.26.

The ANOV A between the moWldS of the three species shows that there are a highly

significant (P<O.OOI) differences for all the selected elements and particle sizes, except

for zinc, which was significantly difference (P<0.05). The Tukey pair-wise comparison,

performed after the Bartlett test for homogeneity of group variances, indicates that the

differences between Amitermes vitiosus and Tumulitermes pastinator mounds are more

important amongst all the elements and particle sizes than-between Amitermes vitiosus

and Nasutitermes triodiae mounds and between Tumulitermes pastinator and

Nasutitermes triodiae mounds (Table 3.26).

146

TABLE 3.26 Selected elements (mg!lOOg) and particle sizes (%) (minimum,

ElementJ

maximum and mean ± standard deviation) of Amitermes vitiosus, Tumulitermes pastinator and Nasutitermes triodiae mounds sampled at sites 1 to 7, together with the probability of differences (P: Av· Tp-Nt) between the three species and the pairwise comparison probabilities between species (P: Av-Tp, P: Av-Nt and P: Tp-Nt).

Av, Tp and Nt p p p p Particle size

Mean ± SD Av·Tp·Nt Av·Tp Av-Nt Tp-Nt

Min Max l l l l Aluminium 2676 7745 4192 ± 1 1 78 ••• ... ... N

Calcium 1 1 .9 !54 54.5 ± 35.2 ... • •• ... ..

Cobalt 0.20 1.21 0.47 ± 0.24 .. • N N

Copper 0.52 2.43 1.01 ± 0.40 •• • N N

Iron 782 6519 2183 ± 1248 ... ... • ...

Potassium 27.7 956 422 ± 278 ... N N ...

Magnesium 37.8 299 91.8 ± 43 ... ... N ...

Manganese 2.23 I l .S 5.34 ± 2.26 ... ... ... N

Sodium 4.61 42.5 18.2 ± 10.7 .. N .. •

Zinc 0.33 1.97 0.80 ± 0.39 • N N •

Clay 6.0 34.1 20.1 ± 5.7 ... ... ... N

Silt 3.8 3 1 .6 10.9 ± 5.8 ... ... ... N

Fine sand 22.6 55.3 39.0 ± 6.5 ... ... N ...

Coarse sand 9.4 47.2 30.2 ± 8.7 ... ... N •

Abbreviations Av• Amilerm£S viliosu.r; Tp Tumulikrmes pa.rtinator; Nt=Na.ru/1/erme.r triodiae t •: O.Ol<P<0.05; ••: O.OOI<P<O.Ol; •u: P<O.OOI

Table 3.26 also illustrates the vast differences between mound elements and particle sizes

between species and sites. For example. the minimum iron content in the mounds studied

was 782 mg/IOOg and the maximum 6519 mg/IOOg (more than 8 times higher). The

results of the probability of differences (ANOVA) between soils of different sites

indicated highly significant differences (P<O.Ol) for all the variables tested (Tables 3.27-

3.28).

The probability of differences between soil and termite mounds by location and species

is given in Table 3.29 together with the percentage differences between the soil and

mound content

TABLE 3.27 Selected elements (mg/100g) and particle size (%) (mean ± standard deviation) of soil samples (0-!0cm) collected at all the site studied: I to 7.

Element/ Daly River Particle size (Site 1)

n�

Daly River (Site 2)

n�

Daly River {Site 3)

n=S

Daly River (Site 4)

n�

Elliott (Site 5)

n�

Howard Springs {Site 6)

n�

Berrimah (Site 7)

n=3

Aluminium 1708 ± 95 2848 ± 208 3187 ± 742 2548 ± 535 2059 ± 389 4544 ± 713 1883 ± 722

Calcium 7.70 ± 3.14- 6.9 ± 3.9 9.6 ± 3.8 12.1 ± 1 1 .7 51.1 ± 12.5 56.8 ± 31.1 17.2 ± 7.2

Cobalt 0.21 ± 0.03 0.44 ± 0.03 0.45 ± 0.06 0.26 ± 0.04 0.28 ± 0.03 0.66 ± 0.25 0.26 ± 0.02

Copper 0.45 ± 0.11 0.66 ± 0.05 0.64 ± 0.1 1 0.55 ± 0.13 0.71 ± 0.06 1.28 ± 0.18 0.59 ± 0.29

Iron

Potassium

1028 ± 57

377 ± 38

1 149 ± 75

389 ± 13

Magnesium 41.4 ± 19.9 88.8 ± 7,3

2809 ± 236

694 ± 158

145 ± 49

801 ± 218 1231 ± 142 6021 ± 956 1442 ± 1543

529 ± 104 103 ± I I

59.6 ± 13.8 53.5 ± 7.4

32.0 ± 12.3 223 ± 61

37.5 ± 6.1 26.0 ± 7.4

Manganese 4.60 ± 3.17 3.08 ± 0.73 7.01 ± 1.92 2.59 ± 1 .06 7.27 ± 3.69 9.04 ± 3.69 2.04 ± 0.08

Sodium 16.8 ± 2.2 9.18 ± 0.40 12.6 ± 1.6 19.4 ± 3.4 4.11 ± 0.20 3.99 ± 0.27 13.6 ± 4.43

Zinc 0.41 ± 0.03 0.72 ± 0.07 1 .50 ± 0.56 0.31 ± 0.12 0.69 ± 0.14 1.85 ± 0.92 0.37 ± 0.16

Clay 6.5 ± 0.3

Silt 8.5 ± 1 . 1

Fine sand 43.8 ± 3.7

Coarse sand 41.0 ± 3.4

#: fl"'3

14.2 ± 1.2

1 1 .4 ± 2.1

34.4 ± 1.5

41.3 ± 2.3

13.2 ± 4.0

20.2 ± 6.4

41.0 ± 1.8

29.0 ± 9.2

13.6 ± 2.2

10.4 ± 2.8

46.3 ± 5.2

34.2 ± 7.6

13.0 ± 3.2

5.2 ± 1.2

32.2 ± 5.5

50.5 ± 4.8

18.9 ± 3.4

7.6 ± 2.0

50.9 ± 4.9

26.8 ± 2.7

10.1 ± 2.7

10.6 ± 1.4

42.2 ± 14.1

35.3 ± 7.3 � .. _,

148

The elements mean content of the soil compared to the mounds and the finer particle

sizes (clay and silt) were generally lower (but not necessary statistically) in most of the

soils studied. An example is given in Figure 3.14 for calcium, potassium and sodium.

The finer soil particle fractions were generally higher in the mound while the larger

particle size fractions (fme sand and coarse sand) were usually higher in the soil at any

given site (Table 3.29) (see Figure 3.15 for clay and coarse sand).

TABLE 3.28 Selected elements (mg/lOOg) and particle sizes (%) (minimum, maximum and mean ± standard deviation) of soil (0-lOcm) sampled at sites 1 to 7, together with the probability (P) of differences between soils.

Element/ Sites 1 to 7 p Particle size Min Max Mean ± SD l

Aluminium 1385 5508 2771 ± 1033 •••

Calcium 4.06 69.1 18.1 ± 19.2 •••#

Cobalt 0.19 0.97 0.38 ± 0.17 ...

Copper 0.37 1.51 0.69 ± 0.28 ...

Iron 505 7371 2219 ± 1777 ...

Potassium 24.0 943 385 ± 260 ...

Magnesium 20.5 201 75.7 ± 52 ...

Manganese 1.66 12.9 5.52 ± 3.29 ..

Sodium 3.75 23.6 1 1 .54 ± 5.73 ...

Zinc 0.20 7.73 1.50 ± 0.76 ..

Clay 6.21 22.05 12.9 ± 4.4 ...

Silt 4 . 1 1 27.17 1 1 .8 ± 6.5 ...

Fine sand 26.34 58.37 41.7 ± 7.6 •••

Coarse sand 20.18 54.09 35.7 ± 9.8 ...

t *: O.Ol<P<0.05; U; O.OOI<P<O.Ol ; •••: P<O.OOI

TABLE 3.29 Probability of differences (P) between soil and termite mound by location and species.

Element/ P t P t P t P t P t P t P t P t P t P t P t Particle size Site I Site 1 Site 2 Site 3 Site 3 soil Site 4 Site 4 Site 5 Site 6 Site 6 Site 7

soil - Tp soil - Th soil - Av soil - Tp • Nt soil - Av soil - Nt soil - Av soil - Tp soil - Nt soil - Nt

Aluminium - ... . .. . N . N . .. - ... - ... - ... - ... • • . ..

Calcium N . N . .. - ... - ... . .. . N - ... N • • . ..

Cobalt - ... - ... . N . N . N • • . N . N . N . N . ..

Copper - ... . .. . N . .. . N - ... - ... - ... . N . N . N

Iron - ... . .. . .. . N - ... . .. - ... - ... .. ... N

Potassium - ... . N • • . N • • - ... . .. - ... . N . N . ..

Magnesium - ... - ... . N . N - ... - ... - ... - ... . .. . .. - ...

Manganese N . N . .. N • • . .. . N . N • .. • •

Sodium - ... . N . .. . N . .. - ... . .. - ... - ... - ... - ...

Zinc . .. . N . N N N • • • • - ... .. ... . ..

Clay - ... - ... . N - ... . .. . N - ... . N - ... - ... • •

Silt . N . N • • . N . .. • • N .. N . N N

Fine sand N N N .. ... N ... . N • • . N

Coarse sand • . N N N N .. . N ... ... .. . •

t: N: P>0.05; *: O.Oi<P<0.05; **: O.OOI<P<O.OI; ***: P<O.OOl -: indicates that the mean of the soil content is lower than the mean of the mound content (but not necessarily statistically). � Abbreviations: Tp = Tumulitermes pastinator; Th = Tumulitermes hastilis; A v = Amitermes vitiosus; Nt = Nasutitermes triodiae .... "'

150

1 50 l

8 1 oo J I 0 � ' "' E � "' 50

0

0

1 000

-Ol 0 0 � ' Ol E �

"

BOO

600

400

200

0

40

0, 30 8 �

ci, 20 s � 1 0

0

Site 1 -

Site 5

§ I I I� Site 7 Site6

Tp Th S Av S Tp Nt S Av Nt S Av S Tp Nt S Nt S -

Site 3

Site 4

Site 1 Site 2 Site 7

Tp Tb 5 Av S Tp Nt S Av Nt S Av S Tp Nt S Nt S

Site 7 Site 4 .--

Site 1

Tp Th Av S Tp Nt S Av Nt S Av S Tp Nt S Nt S

SPECIES I SOIL

FIGURE 3.14 Soil�mound effects (mean ± SE) on calcium, potassium and sodium (mg/lOOg) in mounds of different species and soils (0-lOcm) sampled at different locations.

S = Soil; Species abbreviations as indicated previously.

30

20 .. ->"'

0 10

0

60 50 � 0>

0 0 40 -' 0> 5 30 "0 c 20 "' Cf.! 0 10

0

- Site 6

Site 3 Site 4 Sit• ' I� Site 7

T

Site 1 Sitel

T al T T

J.

Ill <=! Ell I �II§

Tp Th S Av S Tp Nt S Av Nt S Av S Tp Nt S Nt S

SPECIES I SOIL

Site 5 Site 1 Site 2

Site 7

Site 6

Tp Th s A• s Tp Nt s A• Nt s A• s Tp No s No s

SPECIES I SOIL

FIGURE 3.15 Soil-mound effects (mean ± SE) on clay and coarse sand (%) in mounds of different species and soils (0-lOcm) sampled at different locations.

S = Soil; Species abbreviations as indicated previously.

151

152

3.3 Hot Water ("Infusion") Extractable Selected Elements from Amitermes

viliosus Mounds (Elliott, Site 5).

Eleven mounds (top and bottom sections), from Elliott (Site 5), were analysed for

selected elements following the method described in chapter 2 (2.5.1). The "infusion"

analyses were performed in duplicate, using two separate samples for each position. The

infusion's results for selected elements were compared to those obtained following acid

(perchloriclnitric acids) extraction (chapter 2, 2.2.4.2) performed on the same mounds

(I I mounds, top and bottom sections).

3.3.1 Comparison of Hot Water ("Infusion") Element Extracts and

Perchloric/Nitric Acid Extracts.

The concentration (mean ± standard) deviation of selected elements of Amitermes

vitiosus mounds and soils (Elliott, Site 5), extracted with hot water ("infusion") and acid

extraction (perchloric/nitric acids), together with the percentage recovery between

extractions, are presented in Table 3.30.

The results show that three of the selected elements were not detected in the "infusion"

extracts: cobalt, copper and zinc; and three other elements: aluminium, iron and

manganese had a very low percentage of recoveries: <0.3 %. The remaining four

elements: calcium, potassium, magnesium and sodium had percentage recoveries of 6.3 7

%, 6.33. %, 2.56 % and 7.31 % respectively. In the soil extracts, the recoveries were

lower for calcium, potassium and magnesium (1 .95 %, 1.84 % and 0.94 % respectively)

but higher for sodium (9 .3 % ).

153

TABLE 3.30 Selected element concentrations (mean ± standard deviation in mgllOOg) following hot water ("infusion'') and acid (perchloric/nitric acids) extractions of eleven Amitermes vitiosus mounds (Elliott, Site 5) and soils, together with the percentage recovery between extractions.

Element Mounds

"Infusion" Acid % n=44 n=22 recovery

Aluminium 0.56 ± 0.59 3664 ± 396 0.02

Calcium 6.69 ± 3.21 105 ± 21 6.37

Cobalt nd 0.30 ± 0.05 nd

Copper nd 0.91 ± 0.08 nd

Iron 0.10 ± 0. 1 1 1738 ± 156 0.01

Potassium 10.3 ± 7.83 162 ± 12 6.33

Magnesium 2.37 ± 1.55 93 ± 7.1 2.56

Manganese 0.02 ± 0.02 7.45 ± 1.96 0.23

Sodium 0.47 ± 0.54 6.49 ± 0.73 7.31

Zinc nd 1.08 ± 0.09 nd nd: not detected; detection limit (mg/JOOg): Co and Zn • 0.02;

3.3.2 Position Effects on Selected Elements

"Infusion" n=2

1.01 ± 0.24

0.92 ± 0.07

nd

nd

0.13 ± 0.01

1.78 ± 0.1 1

0.45 ± 0.03

o.oz ± 0.00

0.38 ± 0.02

nd Cu .. 0.01

Soil

Acid n=2

1847 ± 330

47 ± 1.9

0.27 ± 0.01

0.68 ± 0.09

1 154 ± 128

97 ± 12

48 ± 5.9

6.37 ± 0.66

4.03 ± 0.01

0.59 ± 0. 1 1

% recovery

0.05

1.95

nd

nd

O.oJ

1 .84

0.94

0.27

9.34

nd

The comparison between the two types of extractions with respect to position (top and

bottom), using ANOVA, is presented in Table 3.31. A significant increase in calcium

in the top section of the mounds is observed following the two types of extractions. A

highly significant increase in iron in the bottom of mounds after hot water extraction

may be attributed to the very low iron concentrations.

154

TABLE 3.31 Position effects on selected elements (mean ± standard deviation in mg/lOOg) in eleven Amitermes vitiosus mounds (Elliott, Site 5) following two types of extraction: hot water ("infusion") extraction and perchloric/nitric acids extraction. ANOV A probability of differences (P) between positions.

Mineral

Top n""22

"Infusion"

Bottom n=22

p �

Aluminium 0.39 ± 0.47 0.73 ± 0.64 N

Calcium 8.06 ± 3.42 5.33 ± 2.35 ••

Cobalt nd nd nd

Copper nd nd nd

Iron 0.05 ± O.o7 0.16 ± 0.1 1 •••

Potassium 12.2 ± 9.09 8.34 ± 5.93 N

Magnesium 2.74 ± 1.27 2.01 ± 1.75 N

Manganese 0.02 ± 0.03 0.02 ± 0.02 N

Sodium 0.62 ± 0.70 0.33 ± 0.24 N

Zinc nd nd nd

t: N: P>O.OS; *: O.Oi<P<0.05; ••: O.OOI<P<O.Ol ; •••: P<O.OOI nd: not detecll:d; detection limit (mg/IOOg): Co and Zn:0.02; Cu: 0.01

Perchloric/nitric extraction

Top n=11

Bottom n=l l

3829 ± 392 3526 ± 359

1 16 ± 17 95 ± 20

0.31 ± 0.05 0.30 ± 0.05

0.94 ± 0.08 0.89 ± O.o7

1 775 ± 160 1701 ± 141

167 ± 12 159 ± 12

97 ± 6.1 89 ± 6.1

7.77 ± 2.01 7.21 ± 1.89

6.67 ± 0.80 6.35 ± 0.63

1 . 12 ± 0.08 1.04 ± 0.08

p � N

•

N

N

N

N •

N

N •

3.4 Soluble Iron, Ionisable Iron and Selected Element Concentrations of

Termitaria and Soils Following Pepsin-Hydrochloric Acid Incubation.

The soluble iron, ionisable iron together with the elements AI, Ca, Co, Cu, K, Mg, Mn,

Na and Zn analyses were performed in triplicate on selected mound and soil samples

(Table 2.1 1) at each site (I to 6) according to the method described in chapter 2.6.2.

3.4.1 Pepsin Concentration Effects on Soluble Iron, Ionisable Iron and Selected

Element content following Pepsin-HCI Acid Incubation.

The results of variation of pepsin concentration (0-0.5 %) in the pepsin-HCl incubation,

on soluble iron, ionisable iron and selected element concentrations (mg/l OOg), together

with the probability of differences between pepsin concentration, are given in Table

3.32.

•

TABLE 3.32 Selected element composition (mg/IOOg) of termitaria reference material® (Nt26D4), following O.IN HCI acid (pH 1.35) extraction with different pepsin percentage v/w (0%, 0.1 o/o and O.So/o), together with the probability P(t) of differences between pepsin concentrations.

Element Pepsin-HCI pH 1.35 extract

0% 0.1% 0.5% P(t)

Aluminium 20.3 ± 0.9 19.8 ± 0.9 19.7 ± 0.9 N

Calcium 33.6 ± 1.0 33.7 ± 0.7 33.7 ± 0.4 N

Cobalt 0.01 ± 0.01 0.01 ± 0.00 O.ot ± 0.00 N

Copper 0.08 ± 0.01 0.09 ± 0.02 0.08 ± 0.01 N

Iron 1 1 .5 ± 0.2 1 1.3 ± 0.3 1 1.2 ± 0.3 N

Potassium 65.8 ± 3.1 65.5 ± 2.9 64.4 ± 1.9 N

Magnesium 44.4 ± 1.4 44.0 ± 0.7 43.6 ± 1.3 N

Manganese 2.39 ± 0.10 2.35 ± 0.09 2.32 ± 0.08 N

Sodium 7.65 ± 0.41 7.43 ± 0.28 7.34 ± 0.24 N

Zinc 0.08 ± O.ot 0.09 ± 0.02 0.08 ± 0.02 N

Iron(II) 8.12 ± 0.27 8.00 ± 0.32 8.04 ± 0.33 N

@: for tennitana material number explanations see chapter 2.1.3. t: N: P>0.05; •: 0.01 <P<0.05 nd: not detected; cobalt and zinc detection limit: 0.02 mgfl OOg.

0%

2.05 ± 0.53

28.7 ± 1.2

nd

0.03 ± 0.00

126 ± 0.27

63.3 ± 1.8

38.2 ± 1.2

1 .89 ± 0.07

0.02 ± 0.02

0.34 ± 0.15

pH 7.5 filtrate

0.1% 0.5%

2.49 ± 0.58 2.90 ± 0.61

28.1 ± 1.6 27.8 ± 1.3

nd nd

0.03 ± O.ot 0.03 ± 0.00

1.49 ± 0.37 1.69 ± 0.37

62.7 ± 2.3 62.4 ± 2.0

37.2 ± 2.2 35.8 ± 2.1

1.82 ± 0.10 1.77 ± 0.12

0.02 ± 0.01 nd

0.38 ± 0.12 0.67 ± 0.92

P())

•

N

N

•

N

•

N

N

� "' "'

156

25.0

� 200 � ' _§ 15.0

f--ctl 1 0 0

2 w _j w 5.0

Pepsjn-HCI Extract fpH 1.35)

0.0 _c_ __ _ 0.0 0. 1

% PEPSIN

§ Aluminium

ISl Soluble iron

� lonisable iron

0.5

FIGURE 3.16 · Pepsin concentration effects on aluminium, soluble iron and ionisable iron (mean ± SE in mg/lOOg) in Nasutitermes triodiae mound (Nt26D4, Daly River, Site 4) following Pepsin-HCI pH 1.35 extraction (n=9).

3.0 pH 7.5 Filtrate

� 2 0 � ' L � Aluminium I � ISl Soluble iron

!:11 lonisable iron 5 f--z w 1.0 2 w _j w

0.0 0.0 0.1 0.5

% PEPSIN

FIGURE 3.17 Pepsin concentration effects on aluminium, soluble iron and ionisable iron (mean ± SE in mg/IOOg) in Nasutitermes triodiae mound (Nt2604) in pH 7.5 filtrates (n=9).

I

I ,

157

The pepsin concentration does not significantly (P>0.05) affect the amount of soluble

iron, ionisable iron and selected elements present in pepsin-HCl acid (pH 1.35) extracts

(see Table 3.32 and Figure 3.!6). In pH 7.5 filtrates. there is a significant

(0.01 <P<0.05) increase in alwninium and soluble iron for the 0.5 % pepsin concentration

(see Figure 3.17) and a significant decrease in magnesium for the same pepsin

concentration. Although the ionisable iron concentration mean is higher in the pH 7.5

filtrate from the 0.5 % pepsin-HCl extract, there is no significant difference. Likewise,

no significant difference has been observed in the other selected elements. In view of

the results obtained (significantly higher soluble iron concentration and higher ionisable

iron mean) a 0.5 % w/v solution of pepsin was used for all incubations as in the in vitro

test for predicting the bioavailability of iron in foods122•

3.4.2 Quality Assurance

The results of the quality assurance are shown in Table 3.33. The internal reference

sample (Nt26D4) was run in duplicate with every batch of samples. The precision of

the method is indicated by the selected element standard deviations (n=40) for the internal reference sample mean contents. From the means of the pepsin-HCl extracts,

the standard deviations are below 5 % for AI, Ca, Fe, K, Mg, Mn, Na, Fe(II). The mean

concentrations of Co, Cu and Zn are very low, 0.02 to 0.07 mg/IOOg and their standard

deviations were 50 %, 14 % and 40 % respectively. The standard deviations from the

mean concentration of selected elements (including soluble iron and ionisable iron) of

pH 7.5 filtrates are much higher.

158

TABLE 3.33 Internal quality control of selected elemental composition (mgllOOg) of pepsin-HCI acid (pH 1.35) extracts and pH 7.5 filtrates of termitaria sample (Nt26D4).

Element Internal reference material: Nt26D4® n=40

Pepsin-HCI

Aluminium 20.6 ± 0.8

Calcium 34.3 ± 1.6

Cobalt 0.02 ± 0.01

Copper O.D7 ± 0.01

Iron 1 1.3 ± 0.5

Potassium 64.1 ± 1.4

Magnesium 46.2 ± 1.3

Manganese 2.39 ± 0.06

Sodiom 7.42 ± 0.31

Zinc 0.05 ± 0.02

Fe(ll) 9.1 7 ± 0.41

pH 7.5 filtrate

2.36 ± 1.12

31 .0 ± 1.8

<0.02

0.03 ± 0.01

1.01 ± 0.35

61 . 1 ± 4.4

42.4 ± 2.6

1.77 ± 0.28

<0.02

0.30 ± 0.1 1 @:lor explanatiOn of temutana reference matenill number reler to chapter :t.J.I

159

3.4.3 Soluble Iron, lonisable Iron and Selected Element Composition of Pepsin

HCI Acid (pH 1.35) Extracts and pH 7.5 Filtrates.

The concentration mean (± standard deviation) of selected elements of Amitermes

viliosus, Tumulitermes pastinator, Tumulitermes hastilis and Nasutitermes triodiae

mounds and soils at different sites (I to 6), in 0.5 % (w/v) pepsin - O.IN HCI acid

(pH 1.35) extracts and pH 7.50 filtrates are given in Tables 3.34 to 3.40, together with

the concentration mean (± standard deviation) of selected elements following

perchloric/nitric acid (4:1) extraction and the percentage recoveries between treatments.

Due to the inherent large degree of variability associated with the exchangeable method

used (O.lN HCl) and the small number of replicates, comparative data between different

species at the same site and same species at different sites have not been statistically

presented, as nearly all comparisons showed no significant differences. Instead,

graphical comparisons are given in Appendices Vl and VII.

3.4.3.1 Soluble Iron, Ionisable Iron and Selected Element Comparisons of

Pepsin-HCI Acid pH 1.35 Extracts, pH 7.5 Filtrates and

Perchloric/Nitric Acid Extracts.

A) Soluble Iron and Ionisable Iron

The concentration mean (± standard deviation) ofperchloric/nitric extractable iron, total

soluble and ionisable iron following pepsin-HCI acid incubation, together with the

percentage of ionisable iron in soluble iron are given in Table 3.34.

As seen in Table 3.34, very little of the iron present in the perchloric/nitric extracts is

released following pepsin-HCl (pH 1.35) incubation. The percentage of soluble iron in

the pepsin-HCl (pH 1 .35) extracts compared to the perchloric/nitric iron varied in

mounds from 0.17 % in Tumu/itermes pastinator at site 6 to 1 1 % in Amitermes vitiosus

at site 4 and in soils from 0.10 % at site 6 to 3.2 % at site 4.

TABLE 3.34 Soluble iron and ionisablc iron content (mean± standard deviation in mg/lOOg) oftermitaria and soils, in perchloric/nitric extracts, pepsin-hydrochloric (pH 1.35) extracts and pH 7.5 filtrates together with the percentage recovery between

Site

Site 1

Site 2

Site 3

Site 4

Site 5

Site 6

ionisablc iron and soluble iron in pepsin-HCI extracts and pH 7.5 filtrates. Depth= I, n=3 unless indicated

Sample Perchloric/nitric Pepsin-HCI (pH 1.35) pH 7.50 filtrate

Iron Soluble iron lron(Il) % Soluble Iron Iron(Il )

Tumulitermes pastinator 1362 ± 150 13.2 ± 3.4 7.42 ± 2.65 56 0.13 ± 0.09 nd

Tumulitermes hastilis 1321 ± 76 18.8 ± 3.9 16.5 ± 3.6 88 2.07 ± 0.71 0.54 ± O.Q2

Soil n=2 1014 ± 80 6.54 ± J .:i9 2.1 I ± 0.18 32 nd nd

Amitermes vitiosus 1497 ± 251 I 13 ± 45 62.5 ± 22.9 55 0.90 ± 0.66 nd

Soil n=2 1 192 ± I I 25.6 ± 13.1 4.19 ± 1 . 19 16 nd nd

Tumulitermes paslinator 2815 ± 162 18.0 ± 10.0 12.9 ± 6.65 72 nd nd

Nasutitermes triodiae 3380 ± 1 1 0 17.0 ± 5.0 14.1 ± 3.8 83 0.36 ± 0.38 0.14 ± 0.17

Soil n=2 2594 ± 350 23.3 ± 13.2 6.10 ± 1.57 26 nd nd Amitermes vitiosus 1368 ± 267 157 ± 77 74.7 ± 80.6 48 4.48 ± 4.17 0.47 ± 0.43 Nasutitermes triodiae 1539 ± 222 15.6 ± 4.8 12.2 ± 4.4 78 5.21 ± 5.13 0.58 ± 0.28 Nt depth=O (old) 1645 ± 184 15.7 ± 4.0 12.4 ± 3.8 79 0.44 ± 0.42 0.26 ± 0.23 Nt deptlv:=O (new) 1 163 ± 73 28.1 ± 3.7 19.5 ± 3.8 69 2.24 ± 1.96 0.48 ± 0. 1 1 Soil n=2 779 ± 1 1 0 24.6 ± 12.6 6.48 ± 1.63 26 nd nd Amilermes vitiosus 1831 ± 279 18.2 ± 2.5 13.1 ± 2.4 72 0.66 ± 0.74 0.21 ± 0.20 Soil n=2 1 1 53 ± 128 5.33 ± 0.54 0.84 ± 0. I I 16 nd nd Tunwlitermes pastinator 5195 ± 561 8.68 ± 1.43 6.22 ± 2.96 72 O.QJ ± 0.05 nd Nasutitermes triodiae 4044 ± 1086 14.9 ± 1.5 13.7 ± 1.4 92 0.34 ± 0.43 0.08 ± 0.14 Soil n=2 6243 ± 1596 6.25 ± 1.29 3.86 ± 0.47 62 nd nd

Abbreviations: nd: not detected; detection limit in mg/IOOg: ' 0.06; iron(ll) 0.20 ; Nt:Nasutitermes triodiae I fOil

%

26

39

I I I I

59 21

32

24

� "' Q

161

The highest ionisable iron contents in pepsin-HCI extracts were found in mounds where

the concentrations ranged from 6.22 mg/1 DOg in Tumulitermes pastinator mounds at site

6 to 74.7 mg/lOOg in Amitermes vitiosus mounds at site 4; while in the soil samples the

ionisable iron ranged from 0.84 mg/JOOg at site 5 to 6.48 mgiiOOg at site 4. In either

case (soluble and ionisable iron) the highest content was found in Amitermes vitiosus

mounds at sites 2 and 4.

As the pH increased from 1.35 in the pepsin-HCl extracts to 7.5 in the filtrates, both the

ionisable and soluble iron decreased. However, the decrease in the ionisable iron was

of a greater magnitude. The ionisable iron contents were very low, ranging in the

mounds, from 0.08 to 0.58 mg/lOOg. Even in Amitermes vitiosus mounds at site 4,

where previously both soluble and ionisable iron were the highest, in pH 7.5 filtrates,

the level was comparable to the other species' mound levels. In soils, at all sites, no

soluble iron nor ionisable iron was detected in pH 7.5 filtrates (for example, soil at site

4: see Table 3.34 and Figure 3.18).

162

1 00

80

f-z 60 w 0 8i 40 Q_

20

perchlortc/njtdc

I f'SW Rr,. 0 Alt"'e �• "' "' • A

1 00

Extraction Procedures: Pepsjn-HCJ

pH 1.35 pH 7 5 filtrate

PerchloricJnjtric Pepsin-HC! pH 1 35

pH 7.5 filtrate 80

f-z 60 w 0 8i 40 Q_

B

20

Q AI CaFe KMg

1 00 .

AI CaFe KMg /\I CaFe KMg

� Aluminium

• Calcium

0 Iron

bl Potassium

113 Magnesium i

Perchlortctnjtric Pepsjn-HCI pH 1.35

pH 7.5 filtrate 80

f-z 60 w 0 8i 40 Q_

20 � Q AJ Ca Fe KMg u � 1.

AI CaFe KMg AI CaFe KMg

C ELEMENT

FIGURE 3.18 Selected element percentage variations between perchloric/nitric acid extracts, pepsinHCI acid pH 1.35 extracts and pH 7.5 filtrates oftennitaria (A: Amitermes vitiosus; B:

Nasutitermes triodiae and soil (C), from Daly River site 4.

TABLE 3o35 Comparison of selected element concentration (mean ± standard deviation in mg/100g) of termite mounds (Tumulitermes pastinator and Tumulitermes hastilis) and soils (0-10cm), sampled from Daly River (site 1) in: A- pepsin-HCI acid (pH 1.35) extracts; 8- pH 7.5 filtrates and C- perchloric/nitric acid (4:1) extracts, together with the % recovery between treatments.

Species Method/ Element ± standard deviation mgllOOg

% recovery Aluminium Calcium Cobalt Copper Iron Potassium Magnesium Manganese Sodium Zinc Iron II

Tp (n 3) A-pH US 29.7 ± .S.9 17.7 ± 5.3 nd 0.04 ± 0.01 13.2 ± 3.4 I 1.0 ± 2.2 8.60 ± 1.28 1.20 ± 0.03 1.84 ± 1.69 0.03 ± 0.04 7.42 ± 2.65

B-pH 7.50 0.71 ± 0.16 16.0 ± 4.7 nd 0.01 ± 0.00 0.13 ± 0.09

% (B/A) 2.4 94 - 13 0.97

C- Total 2959 ± 248 21.5 ± 5.1 0.29 ± 0.01 0.56 ± 0.02 1362 ± 150

% (NC) 1.0 80 - 7.4 1.0

% (B/C) 0.02 74 - 0.96 0.01

IO . .S ± 2.1

95

490 ± 24

2.2

2.1

7.84 ± 1.23 0.79 ± 0.05

91 66

63.1 ± 5.1 3.27 ± 0.12

1 4 37

12 24

0 nd nd

26.8 ± 3.8 0.46 ± 0.03

6.9 7.6

Th (n=3) A-pH 1.35 33.5 ± 3.0 60.5 ± 27.6 0.02 ± 0.02 0.06 ± 0.02 18.8 ± 3.9 16.7 ± 7.7 22.7 ± 9.7 2.97 ± 1.40 1.22 ± 0.62 0.10 ± 0.05 16.5 ± 3.6

B·pH 7.50

% (B/A)

c. Total

% (NC)

% (B/C)

2.88 ± 0.62 53.1 ± 24.0 nd 0.02 ± 0.01

8.6 88 nd 29

2364 ± 180 68.8 ± 30.6 0.32 ± 0.04 0.58 ± 0.10

1.4 88 7.0 9.8

0.12 77 - 2.9

2.07 ± 0.71 16.2 ± 8.39 20.3 ± 9.3 2.04 ± 1.01 -I I 97 90 69 0

1321 ± 76 407 ± 21 68.6 ± 10.0 5.16 ± 1.36 18.9 ± 1.0

1.4 4.1 33 " 6.5

0.16 3.9 30 40

Soil (n=2) A-pH 1.35 32.1 ± 8.2 4.11 ± 1.88 nd 0.01 ± 0.00 6.54 ± 1.29 2.29 ± 0.09 0.52 ± 0.24 0.37 ± 0.14 0.56 ± 0.01

B-pl-1 7.50

% (B/A)

C· Total

% (NC)

% (B/C)

0.62 ± 0.15 4.17 ± 1.89

2.0 102

nd nd

- -

1665 ± 67 8.83 ± 2.63 0.20 ± 0.00 0.38 ± 0.01

1.9 47 - 2.7

0.04 48 - -

nd

-1014 ± 80

0.65

-

Abbreviations: Tp = Tumulitermes pastinator; Th = Tumulitermes hastilis; nd = not detected Detection limit (mg!IOOg): Co and Zn = 0.02; Cu = 0.01; Fe = 0.06; Fe(II) = 0.20

nd 0.67 ± 0.16 0.16 ± 0.05 -- l30 42

348 ± 26 31.2 ± 0.5 3.36 ± 0.24 15.9 ± 2.0

0.66 1.7 I I 3.5

0 2.2 4.6

nd

-0.44 ± 0.08

23

0.54 ± 0.02

3.3

0.06 ± 0.06 2.11 ± 0.18

nd nd

0.40 ± 0.01

1 5

� "' ...

TABLE 3.37 Comparison of selected element concentration (mean ± standard deviation in mg/100g)) of termite mounds (Tumulitermes pastinator and Nasutitermes triodine) and soils (Q-10cm), sampled from Daly River (site 3) in: A- pepsin-HCI add (pH 1.35) extracts; 8- pH 7.50 filtrates and C- perchloric/nitric acid (4:1) extracts together with the % recovery between treatments.

Species Method/ Element ± standard deviation mgllOOg

%recovery Aluminium Calcium Cobalt Copper Iron Potassium Magnesium Manganese Sodium Zinc Iron II

Tp (n 3) A-pH 1.35 29.6 ± 6.8 19.8 ± 5.0 0.04 ± 0.00 0.94 ± 0.52 18.0 ± 10.0 17.4 ± 3.1 29.0 ± 6.4 2.39 ± 0.46 1.78 ± 0.58 0.66 ± 0.23 12.9 ± 6.65

B-pH 7.50 0.99 ± 0.51 19.3 ± 5.1 nd 0.15 ± 0.06 nd 17.9 ± 2.8 27.0 ± 5.9 1.78 ± 0.53 0 nd nd

% (B/A) 3.3 97 16 103 93 74

C- Total 3715 ± 263 21.9 ± 3.0 0.39 ± 0.01 1.40 ± 0.43 2815 ± 162 624 ± 41 145 ± 17 5.43 ± 0.36 12.4 ± 1.7 1.51 ± 0.33

%(A/C) 0.80 91 10 67 0.64 2.8 20 44 14 44

% (B/C) 0.03 88 - II - 2.9 19 33

Nt (n=3) A-pH 1.35 28.5 ± 2.4 22.5 ± 6.4 0.02 ± 0.01 0.20 ± 0.11 17.0 ± 5.0 24.7 ± 0.4 56.8 ± 27.3 2.44 ± 0.95 3.49 ± 2.47 0.16 ± O.G7 14.1 ± 3.8

B-pH 7.50 0.67 ± 0.12 21.2 ± 5.6 0.01 ± 0.02 0.05 ± 0.05 0.36 ± 0.38 16.7 ± 5.6 52.5 ± 28.7 1.75 ± 1.02 nd 0.14 ± 0.17

% (B/A) 2.4 94 56 24 2.1 67 93 12 - 1.0

C- Total 4544 ± 113 23.9 ± :5.86 0.51 ± 0.05 0.71 ± O.ll 3380 ±Ito 864 ± 78 268 ± 27 9.23 ± 1.27 16.4 ± 3.2 1.5:5 ± 0.04

%(A/C) 0.63 95 3.9 28 0.5 2.9 21 26 21 10

% (B/C) 0.01 89 2.2 6.9 0.01 1.9 20 19

Soil (n=2) A-pH 1.35 31.2 ± 5.7 7.93 ± 1.94 0.02 ± 0.03 0.08 ± 0.04 23.3 ± 13.2 7.32 ± 2.88 13.8 ± 7.3 0.85 ± O.ll 1.47 ± 0.42 0,07 ± 0.10 6.10 ± 1.57

B-pH 7.50 0.27 ± 0.02 6.71 ± 1.62 nd nd nd 6.33 ± 3.04 11.7±6.3 0.52 ± 0.16 - nd nd

% (B/A) 0.88 85 - - 81 85 61

C- Total 3801 ± 146 12.0 ± 1.3 0.48 ± 0.05 0.51 ± 0.02 2594 ± 350 853 ± 128 179 ± 30 6.18 ± 0.63 14.2 ± 0.4 1.36 ± 0.38

%(NC) 0.82 66 3.7 15 0.90 0.86 1.1 14 10 5.1

% (B/C) 0.01 56 - 0.74 6.5 8.4

Abbreviations: Tp = Tumulitermes pastinator; Nt = Nasutitermes triodiae; nd = not detected ~

'"' Detection limit (mgllOOg): Co and Zn = 0.02; Cu = 0.01; Fe= 0.06; Fe(ll) = 0.20 "'

-"' TABLE 3.38 Comparison of selected element concentration (mean ± standard deviation in mg/100g) of termite mounds (Nasutitermes triodiae) Q\

and soils (D-10cm), samples from Daly River (site 4), depth"" 0 (new/old material), in: A- pepsin-hydrochloric acid (pH 1.35) extracts; 8- pH 7.50 filtrates and C- perchloric/nitric acid (4:1) extracts together with the% recovery between treatments. Probabilities (P) of differences between ages (new/old).

Species/ Method/ Elementl ± standard deviation mg/IOOg

P()) % recovery Aluminium Calcium Cobalt Copper lro" Potassium Magnesium Manganese Sodium Zinc Iron II

Nt (old) A-pH 1.35 21.1 ± 4.0 25.8 ± 8.5 0.02 ± 0.02 0.07 ± 0.01 15.7 ± 4.0 38.1 ± 12.0 25.8 ± 10.0 1.72 ± 0.78 2.97 ± 0.57 0.08 ± 0.01 12.4 ± 3.8

n=J B-pH 7.50

% (8/A)

C- Total

%(A/C)

% (8/C)

1.18 ± 0.75

5.6

22.4 ± 7.3

87

4393 ± 105 25.9 ± 7.7

0.5 99

0.03 86

"' 0.02 ± 0.01 0.44 ± 0.42

34 2.8

0.31 ± 0.05 0.85 ± 0.03 1645 ± 184

6.1 8.4

2.4

0.95

O.QJ

38.0 ± 11.5

100

623 ± 17

6.1

6.1

22.2 ± 9.0

86

101 ± 17

25

22

0.91 ± 0.32

53

2.11 ± 0.60

81

43

24.2 ± 3.3

12

0.03 ± 0.02 0.26 ± 0.23

33 2.1

0.49 ± 0.04

16

5.2

Nt (new) A-pH 1.35 18.4 ± 3.1 25.3 ± 9.0 0.04 ± 0.01 0.10 ± 0.02 28.1 ± 3.7 38.8 ± 15.7 28.0 ± 3.0 2.31 ± 0.89 4.21 ± 1.18 0.10 ± 0.04 19.5 ± 3.8

n=3 B-pH 7.50 1.70 ± 1.50

%(8/A) 9.2

C- Total

%(A/C)

% (8/C)

PU) A- pepsin-HCI

P(t) 8- pH 7.50 filtrates

PU) C- perchloriclnitric

3382 ± 85

0.54

0.05

N

N ...

21.9 ± 7.5

87

26.6 ± 8.4

95

82

N

N

N

"' "' 0.27 ± 0.04

14

N

N

0.04 ± 0.01 2.24 ± 1.96 39.2 ± 16.5

40 8.1

0.69 ± 0.04 t 163 ± 73

14 2.4

5.6

N

N .. 0.19

• N

•

101

579 ± 45

6.7

6.8

N

N

N

25.1 ± 2.6

90

91.0 ± 6.5

31

28

N

N

N

1.46 ± 0.59

63

2.58 ± 0.64

90

57

N

N

N

Abbreviations: Nt = Nasutitermes triodiae; nd = not detected; P = probability of differences between material ages (new/old) Detection limit (mg/lOOg): Co and Zn = 0.02; Cu = 0.01; Fe(ll) = 0.20 P(t): N: P>0.05; *: 0.01< P<O.OS; u: O.OOI<P<O.Ol; ***: P<O.OOI

21.5 ± 5.4

20

N

N

0.02 ± 0.02 0.48 ± 0.11

15 25

0.43 ± 0.04

24

3.7

N

N

N

N (0.083)

N (0.202)

TABLE 3.39 Comparison of selected element concentration (mean ± standard deviation in mg/100g) of termite mounds {Amitennes vitiosus and Nasutitermes triodiae) and soils ({).10cm), samples from Daly River (site 4), in: A· pepsin-HCI acid (pH 1.35) extracts; 8- pH 7.50 filtrates and C- perchloric/nitric acid (4:1) extracts, together with the% recovery between treatments.

Species Method/ Element ± standard deviation mg/1 OOg

%recovery Aluminium Calcium Cobalt Copper !roo Potassium Magnesium Manganese Sodium Zinc Iron II

Av (n-3) A-pH 1.35 49.5 ± 26.1 52.9 ± 59.3 O.o9 ± O.o7 0.15 ± 0.08 157 ± 77 16.5 ± 12.3 33.4 ± 27.1 3.85 ± 3.92 4.17 ± 1.28 0.11 ± 0.12 74.7 ± 80.6

B-pH 7.50 1.31 ± 0.94 38.9 ± 42.0 0.01 ± 0.02 0.04 ± O.oJ 4.48 ± 4.17 15.5 ± 11.8 28.3 ± 22.3 1.50 ± 1.47 od 0.47 ± 0.43

% (BIA) 2.7 74 " 27 2.9 93 84 39 0.63

C- Total 3954 ± 514 58.4 ± 64.6 0.41 ± 0.10 0.84 ± 0.17 1368 ± 267 754 ± 138 Ill ± 45 5.19 ± 3.44 34.0 ± 8.4 0.46 ± 0.15

%(A/C) 1.3 91 22 18 II 2.2 30 74 12 25

% (B/C) 0.03 67 3.3 4.7 0.33 2.1 26 29

Nt (n="4) A-pH 1.35 19.6 ± 3.5 36.2 ± 7.5 0.02 ± 0.01 0.09 ± 0.02 15.6 ± 4.8 54.9 ± 7.9 46.7 ± 16.4 2.45 ± 0.57 9.33 ± 5.10 0.07 ± 0.02 12.2 ± 4.4

B-pi I 7.50 6.05 ± 5.31 33.2 ± 7.5 0.01 ± 0.01 0.05 ± 0.02 5.21 ± 5.13 54.1 ± 8.1 43.2 ± 15.2 1.97 ± 0.45 . 0.01 ± 0.02 0.58 ± 0.28

% (B/A) 31 92 77 58 33 99 93 81 . 12 4.8

C- Total 4109 ± 573 38.5 ± 7.4 0.33 ± 0.02 0.85 ± 0.05 1539 ± 222 687 ± 44 114 ± 19 3.66 ± 0.52 31.6 ± 3.9 0.42 ± O.o4

%(A/C) 0.48 94 5.0 10 1.0 8.0 41 67 30 16

% (B/C) 0.15 86 3.9 6.0 0.34 7.9 38 54 2.0

Soil (n=2) A-pH 1.35 37.0 ± 14.7 2.58 ± 0.08 0.02 ± 0.00 0.07 ± 0.04 24.6 ± 12.6 5.85 ± 1.86 3.83 ± 0.86 0.14 ± 0.01 1.47 ± 0.16 0.04 ± 0.01 6.48 ± 1.63

B-pH 7.50 od 2.09 ± 0.26 od od od 5.73 ± 1.79 3.01 ± 0.78 0.03 ± 0.01 . od od

% (B/A) 81 98 79 22

C- Total 2657 ± 779 6.07 ± 0.63 0.25 ± 0.04 0.58 ± 0.18 779 ± 110 551 ± 144 57.4 ± 13.9 2.30 ± 0.08 20.3 ± 4.7 0.32 ± 0.14

%(A/C) 1.4 42 7.3 12 3.2 1.06 6.7 6.2 7.3 II

% (8/C) 0.01 34 . 1.6 5.2 1.4

Abbrev~atlons: A' Amitermes vmosus; Nt Nasu/ltermes tridwe; od not detected Detection limit (mg/100g): Co and Zn = 0.02; Cu = 0.01; Fe= 0.06; Fe(ll) = 0.20 ~

"' ....

TABlE 3.40 Comparison of selected element concentration (mean ±standard deviation in mg/100g) of termite mounds (Tumulitermes pastinator ~ and Nasutitermes triodiae) and soils (D-10cm), samples from Howard Springs (site 6), in: A- pepsin-HCI (pH 1.35) extracts;: 8- pH 7.50 QO

filtrate and C- perchloric/nitric acid (4:1) extracts, together with the% recovery between treatments.

Species Method/ Element ± standard deviation mg/IOOg

%recovery Aluminium Calcium Cobalt Copper Iron Potassium Magnesium Manganese Sodium Zinc Iron II

Tp (n-3) A-pH 1.35 44.2 ± 10.2 36.5 ± 20.4 0.02 ± 0.03 0.14 ± 0.08 8.68 ± 1.43 5.02 ± 2.27 11.6 ± 5.5 1.50 ± 1.08 0.91 ± 0.38 O.o? ± 0.04 6.22 ± 2.96

B-pH 7.50 0.96 ± 0.37 35.1 ± 19.7 "' 0.03 ± 0.02 O.o3 ± 0.05 4.24 ± 2.62 11.3 ± 5.7 1.03 ± 0.71 - "' "' % (B/A) 2.2 96 - 19 0.35 84 98 68 C- Total 6557 ± 1029 40.8 ± 19.1 0.80 ± 0.26 1.76 ± 0.59 51~5 ± 561 40.8 ± 11.7 48.5 ± 7.5 5.54 ± 1.90 6.20 ± 1.38 1.12 ± 0.30

%(NC) 0.67 89 2.7 8.0 0.17 12 24 27 15 6.2

% (B/C) 0.01 86 1.5 0.01 10 23 19

Nt (n"'3) A-pll 1.35 48.4 ± 4.3 74.6 ± 20.0 0.03 ± 0.05 0.14 ± 0.02 14.9 ± 1.5 12.4 ± 3.5 19.4 ± 5.4 2.08 ± 1.15 1.58 ± 0.51 0.22 ± 0.04 13.7 ± 1.4

B-pH 7.50 1.53 ± 0.72 69.7 ± 16.7 0.01 ± 0.01 0.05 ± 0.01 0.34 ± 0.43 12.1 ± 3.5 17.8 ± 4.5 1.36 ± 0.73 "' 0.08 ± 0.14

% (B/A) 3.2 94 20 33 2.3 98 92 65 0.57

C- Total 5579 ± 692 83.5 ± 21.0 0.73 ± 0.20 1.48 ± 0.12 4044 ± 1086 37.1 ± 3.6 48.9 ± 8.4 5.03 ± 2.46 5.30 ± 0.54 1.01 ± 0.11

%(NC) 0.87 89 3.8 9.3 0.37 33 40 42 30 22

% (B/C) O.Q3 84 0.8 3.1 0.01 33 37 27

Soil (n=2) A-pH 1.35 57.8 ± 16.2 37.6 ± 8.6 0.03 ± 0.04 0.10 ± 0.01 6.25 ± 1.29 3.46 ± 2.21 6.75 ± 1.42 1.67 ± 0.71 1.15 ± O.Q9 1.20 ± 1.01 3.86 ± 0.47

B-pH 7.50 0.91 ± 0.01 33.5 ± 8.0 0.01 ± 0.02 "' "' 2.48 ± 1.07 5.46 ± 1.12 1.00 ± 0.52 "' "' % (8/A) 1.6 89 45 72 81 60

C- Total 4097 ± 436 45.5 ± 11.4 0.60 ± 0.23 1.26 ± 0.06 6243 ± 1596 37.9 ± 17.5 33.9 ± 3.9 8.50 ± 4.25 3.94 ± 0.23 1.87 ± 1.16

%(NC) 1.4 83 5.0 7.6 0.10 9.1 20 20 29 64

% (B/C) 0.02 74 2.2 - - 6.5 16 12

Abbreviations: Tp = Tumulitermer partinator; Nt = Narutitermer triodiae; nd = not detected Detection limit (mg/IOOg): Co and Zn = 0.02; Cu = 0.01; Fe= 0.06; Fe(II) = 0.20

169

B) Selected Elements

As Tables 3.35 to 3.40 show, very little of the predominant elements (aluminiwn and

iron) present in the mounds and soil, at all sites, were released with pepsin-HCI acid at

pH 1.35. The percentage of recoveries of aluminium in the pepsin-HCI acid extracts

compared to the perchloric/nitric acid extracts were very low, less than 2 % for all

species, at all sites. The highest recoveries in the pepsin-HCl extracts of mounds,

compared to the perchloric/nitric acid extracts, were for calcium (greater than 78 % for

all species, at all sites).

The recoveries varied with respect to the origin of the sample. Generally, the percentage

recoveries with pepsin-HCl extracts for calcium, potassium, magnesium, manganese and

sodium were lower in the soil. This trend was not observed between mounds and soils

sampled in Elliott, where the percentage recoveries were similar for calcium, potassium,

magnesiwn and sodiwn (Table 3.36).

When the pH was increased from 1.35 to 7.5, the amount of aluminium, cobalt, copper,

soluble iron, ionisable iron and zinc decreased dramatically for all species, at all sites.

The cobalt, copper and zinc contents in the pH 7.5 filtrates were close to the detection

limit or not detected at all. Very little of the calcium, potassium and magnesium were

lost after neutralisation to pH 7.5, the percentage recoveries of these elements remained

between 80-100% for all species and at all sites (Table 3.35 to 3.40). The manganese

recoveries in the pH 7.5 filtrates compared to pepsin-HCl extracts were more variable

with an average of 65 % for all species termitaria and 49 % for all soils. Here again

as in the pepsin-HCl extracts, the percentage recoveries for calcium, potassium,

magnesium and manganese were generally lower in the soil than in the mounds, as

shown in Figure 3.19 for Nasutitermes triodiae, Tumulitermes pastinator and soil from

site 3.

170

100 D Nt

80 j ~ I bl Tp >- Ill! Soil a: w 60 > 0 () w 40 a:

"' 20

0 Calcium PotaSSIUm Magnesium Manganese

ELEMENT

FIGURE 3.19 Selected elements (calcium, potassium, magnesium and manganese) percentage recovery in pH 7.5 filtrates of Nasutitermes triodiae (Nt) mounds, Tumulitermes pastinaror (Tp) mounds and soils·at site 3.

171

Figure 3.18 shows an example of the change in the percentage distribution of aluminium,

calcium, iron, potassium and magnesium in perchloric/nitric extracts, pepsin-HCl extracts

and pH 7.5 filtrates. At all sites and for all species, the aluminium and iron were the

dominant elements in the perchloric/nitric extracts. In the pH 7.5 filtrates, the 3

dominant elements were calcium, potassium and magnesium. Their relative percentages

varied according to the type of samples. For example, in the pH 7.5 filtrates, at site 4

calcium is the dominant element in Amitermes vitiosus mounds while potassium is the

dominant element in Nasutitermes triodiae mounds and soils.

3.4.3.2 Age Effects on Selected Elements (Depth=O).

As shown in Table 3.38, the results of the AN OVA between ages of mound material

(old and new), indicate a significant increase (0.01 <P<0.05) in the new material for

soluble iron in the pepsin-HCl (pH 1.35) extract and a significant decrease for iron

following the perchloric/nitric acid extraction. A smali ionisable iron increase (P=0.083)

in the new material was detected in the pepsin-HCI extract. Although highly significant

decreases were observed in new material for aluminium and copper in the

perchloric/nitric extracts, no other differences were observed in the pepsin-HCl extracts

and pH 7.5 filtrates.

3.4.3.3 General Oveniew of Different Species Studied at Different Sites with

Relation to the Adjacent Soil (0-lOcm).

The minimum, maximum and mean ± standard deviation of selected element contents

(mg/1 OOg) of the species of mounds. chosen by the Aboriginal communities, and the

adjacent soils (0-lOcm) sampled at sites I to 6, in pepsin-HCl extracts and pH 7.5

filtrates are given in Tables 3.41 and 3.42 respectively.

172

TABLE 3.41 Selected elements (mg/100g) (minimum, maximum and mean ± standard deviation) of Amitermes vitiosus, Tumulitermes pastinator and Nasutitermes triodiae mounds and soils (O~lOcm) sampled at sites 1 to 6, following pepsin-HCl incubation (pH 1.35).

Element Av, Tp and Nt (n=28) Sites 1 to 6 (n==12)

Min Max Mean ± SD Min Max Mean ± SD

Aluminium 15.3 79.2 35.6 ± 15.6 9.50 69.3 36.5 ± 17.3

Calcium ll.S 128 44.4 ± 36.7 1.74 50.4 17.4 ± 19.3

Cobalt nd 0.17 0.04 ± 0.04 nd 0.06 0.02 ± 0.02

Copper O.o3 1.45 0.22 ± 0.30 0.01 0.12 0.07 ± 0.04

Soluble iron 7.33 245 40.7 ± 56.7 4.95 34.8 15.3 ± 11.8

Potassium 3.44 62.3 20.3 ± 16.1 1.90 10.4 5.47 ± 3.05

Magnesium 5.25 88.2 27.7 ± 20.4 0.35 18.9 6.45 ± 5.60

Manganese 0.50 8.38 2.65 ± 1.69 0.13 2.26 0.94 ± 0.76

Sodium 0.51 16.9 3.25 ± 3.35 0.26 1.77 1.05 ± 0.44

Zinc nd 0.82 0.17 ± 0.20 nd 1.91 0.23 ± 0.55

Ionisable iron 4.30 167 23.7 ± 33.4 0.76 7.63 3.93 ± 2.24

Abbreviations: Av= Amitermes vitiosus; Tp= Tumulitermes pastinator; Nt= Nasutitermes triodiae nd= not detected; detection limit (mgflOOg): Co and Zn c 0.02

The selected element content differences (%) between the soil and the termite mounds

at a given site in pepsin-HCI extracts and pH 7.5 filtrates are given in Table 3.43. In

pepsin-HCI extracts, the element mean contents of the soil compared to the mounds were

generally lower in most of the soils studied, with the exception of aluminium where they

were higher in some cases. In pH 7.5 filtrates, the element mean contents of the soil

compared to the mounds, when detected, were always lower.

173

TABLE 3.42 Selected element content (mgllOOg) (minimum, maximum and mean ± standard deviation) ofAmitermes vitiosus, Tumulitermes pastinator and Nasutitermes triodiae mounds and soils (0-lOcm) sampled at sites 1 to 6, in pH 7.5 filtrates.

Element Av. Tp and Nt (n-28) Sites 1 to 6 (n-12)

Min Max Mean ± SD Min Max Mean ± SD

Aluminium 0.37 13.9 1.72 ± 2.57 nd 0.92 0.48 ± 0.35

Calcium 10.8 119 39.9 ± 32.0 1.54 45.7 15.7 ± 17.6

Cobalt nd 0.05 nd nd 0.03 nd

Copper nd 0.21 0.05 ± 0.05 nd 0.01 nd

Soluble iron nd 12.6 1.48 ± 2.91 nd nd nd

Potassium 2.04 63.1 18.8 ± 16.1 nd 9.50 4.69 ± 3.51

Magnesium 4.59 85.7 25.4 ± 19.0 0.475 16.1 5.47 ± 4.91

Manganese 0.36 4.01 1.64 ± 0.91 0.03 1.68 0.57 ± 0.57

Sodium nd

Zinc nd 0.03 nd nd nd nd

Ionisable iron nd 1.00 0.18 ± 0.28 nd nd nd

Abbreviations Av=- Amitermes vitiosus; Tp= Tumulitermes pastinatar; Nt=Nasutitennes triodwe

nd= not detected; detection limit (mgiiOOg): Co and Zn = 0.02; Cu - 0.01; Fe = 0.06; Fe(! I) .. 0.20

CHAPTER FOUR

DISCUSSION

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175

4 DISCUSSION

4.1 Introduction

Although substantial chemical and physical analyses have been reported98•77

•136 in

Australia. the literature relating to the Aboriginal nutritional/medicinal uses of mounds

is negligible. As indicated in section 1.2.4.3-F only one study136 mentioned iron values

in mounds and this for two Amitermes species in Queensland. Therefore, as described

in section 1.5, the first part of the analyses (nitric/perchloric acid extraction) was to

determine the selected elemental composition of termite mounds eaten by the Aboriginal

communities and the variation between and within mounds together with particle size

analyses. The selection of elements was determined on the basis of the Aboriginal use

of termite mounds (sections 1.1.1 and 1.1.2). The second part of this study, the hot

water "infusion" analyses, reflects the way the Aborigines (Elliott) prepare their

termitaria for nutritional/medicinal purposes. In ·view of the possible

nutritional/medicinal value ascribed to termite mounds by Aboriginal people, the "bio

availability" of elements was determined, with an emphasis on iron bio-availability.

4.2 Acid Extractable (Perchloric/Nitric Acid) Selected Element Concentrations

Together with Particle Size

4.2.1 Quality Control and Quality Assurance

For the purpose of studying the selected elemental composition of termite mounds and

the variation within mounds and between mounds, a method that would give good

p~ecision, so as to allow valid statistical analyses, was selected. As it was not necessary

for the purpose of this study to select a method that would give a total extraction, the

major focus being the human nutrition, a perchloric/nitric acid (4:1) extraction was

selected. Perchloric/ nitric acid mixture is very effective at destroying organic matter

but, as expected, cannot break down certain soil minerals, silicates in particular, so the

total element concentration of certain elements, for example, AI, K, Na. cannot be

176

obtained. In order to determine the quality of our method, the selected element

percentage recovery versus known reference material and internal termitaria reference

material were performed.

The recoveries for the internal reference tennitaria material compared to external

laboratory results show that the perchloriclnitric acid method gives comparable results

to XRF, except for manganese in Tumulitermes pastinator and Nasutitermes triodiae

mounds. The low percentage recovery in this case could be explained by the fact that

the XRF detection limit for manganese as MnO was 0.01 % which translates to a

detection limit in samples of 7.8 mg/IOOg. Being at the detection limit, it is possible

that in the case of Tumulitermes pastinator and Nasutitermes triodiae, the results have

been over-estimated. This hypothesis is supported by the fact that the ICP results for

Tumulitermes pastinator and Nasutitermes triodiae of the external laboratory are much

closer to our results (63 and 66 % respectively). It was expected that the external

laboratory results with XRF and JCP (acid digestion mixture including HF) would be

higher than those from perchloric/nitric extraction. The XRF calcium and magnesium

results (Table 3.2) are much lower than the ICP results and the perchloric!nitric results

(mainly for Tumulitermes pastinator and Nasutitermes triodiae). Here again the values

were close to the XRF calcium and magnesium detection limit which were 0.01 % CaO

or 7.2 mg of calcium per 100 g and 0.05% MgO or 30 mg of magnesium per 100 g.

It is possible that in this case the XRF values have been underestimated. The high K

percentage recovery in Elliott (1 00 % in Amitermes vitiosus mound and 86 % in soil)

would indicate that the potassium is in a different form and is more readily extracted in

Amitermes vitiosus mound and soil in Elliott than in Amitermes vitiosus mounds in Daly

River (Av44D4, Table 3.2). For iron, there is good agreement for all mounds and soil

material between the XRF, ICP (mixed acid digest) and perchloric/ nitric extraction.

This was also observed with the reference material BCSS-1 and MESS-I.

The quality assurance, monitoring the instrumental, procedural and time variations shows

the precision of the method to be very high. This is indicated by the low standard

deviations (usually below 5 %, but forK and Na 10 %) from the selected element mean

following perchloric/nitric extractions (Table 3.2).

177

4.2.2 Overview, General Correlation

As ind.icated in section 3.2.2, a complete study of the correlations between all variables

studied may have been interesting but it was not the focus of this project. In relation

to the Aboriginal uses of termitaria, the general correlation between clay and other

variables is of particular significance. As indicated previously (section 3.2.2), the clay

is positively correlated to aluminium, cobalt, copper and iron and negatively correlated

to potassium, magnesium, silt and coarse sand.

The positive correlation between clay and aluminium, copper and uon could be

explained by the fact that:

the structure of the clay itself is commonly a combination of Al-OH octahedra

(gibbsite sheet) and Si-0 tetrahedra (silica sheet)';

copper exists in soil mainly as the divalent Cu2+ which is adsorbed by clay minerals

or is associated with organic matter•;

iron{III} oxides and oxy-hydroxides of iron are the most abundant non-clay minerals

found in the clay fractions of soils130• They occur as coatings of aggregates or as

separate constituents of the clay fraction94; recent evidence shows that small amounts

of iron {2 %) may replace aluminium in the kaolin structure130•

The negative correlation between potassium and clay may indicate that the type of clay

of the termite mound could mainly be kaolinic as reported by Barber (1984)1'. Soils

whose clays are primarily kaolinic do not fix potassium. This may be reinforced by the

negative correlation between clay and magnesium, as magnesium is not a constituent of

kaolinic soil clays but of soil clays such as montmorillonite, vermiculite, chlorite and

illiteu. The coarse sand and silt are also negatively correlated to clay. This is to be

expected as their percentage compositions are interrelated, with the total particle fraction

(clay + silt + fine sand + coarse sand) being 100 percent. Another factcr is that

although generally the clay particles have developed from weathering of sand and silt

particles, most clay particles differ mineralogically from sand and silt. The

mineralogical composition of sand and silt may be similar11 • This hypothesis (the

mineralogical difference between clay and the sand and silt) is reinforced by the fact that

178

in contrast to the clay, the silt and the coarse sand are positively correlated to potassium

and magnesium (see Table 3.4).

The previously reported soil and mound studies in Northern Australia (section 1.2.3.1)

support the hypothesis that kaolin could be a major part of the clay fraction of the

termitaria and soil in Ibis study. As reported by Norrish and Pickering (1983)130, in

Northern Territory, kaolin is the dominant mineral in clays followed by illite130• The

results of tbe mineralogical analyses conducted by Lee & Wood (!971b)" (Table 1.3),

indicated a high percentage of kaolin type of clay in the termitaria and soils studied in

the Northern Territory frOm Howard Springs to Tennant Creek. The kaolin constituted

more that 80 % of the clay in Howard Springs and Larrimah; in Daly River 65-85 %

kaolin was found inNasutitermes triodiae mound clay. As discussed in section 1.2.3.1,

the kaolin ratio in the subsoil (B horizon) was generally greater than in the topsoil and

tbe clay of half of tbe mounds studied by Lee & Wood (!97lb)" came entirely from

subsoil. This was always the case for Nasutitermes triodiae mounds studied at 3

geographically different locations98• For example, in Daly River the high kaolin

composition of the clay inNasutitermes triodiae mounds came from the B horizon (65-

80 % kaolin); tbe A horizon had only 30-50 % kaolin".

In relation to the Aboriginal use of termite mounds the presence of kaolin type clay

could be of major importance. Kaolin has long been used for the treatment of gastric

disorders in both traditional and modem pharmacologym (section 1.1.4). It has also

been used in the treatment of chronic ulcerative colitis to absorb bacteria and toxins in

the colon and is usually given as a mixture (20: I) with pectin in a sweetened suspension

for the treatment of abnormal intestinal fermentation178• This could be related to the

Aboriginal use of termitaria for upset stomach, diarrhoea, stomach aches and after eating

certain food (section 1.1.2.1).

Another important aspect of the possible high concentration of kaolin in the clay fraction

in the mounds studied is in relation to the low cation-exchange capacity of kaolin clay.

Hence, the reduced possibility of nutritional complications of iron absorption. The

cation-exchange capacity (CEC) is a measure of tbe total of tbe negative charges of tbe

179

clay. Kaolin belongs to the 1:1 type of clay (each sheet has one silica tetrahedral layer

and one alumina octahedral layer). The 1:1 clays have little if any isomorphous

substitution and, hence, low negative chargeu. The cation-exchange capacity of kaolin

and montmorillonite ranges from 1 to 15 and 80 to 150 me/100g11, respectively.

Minnich et a! (1968)114 reported the possible complication of clay ingestion due to iron

absorption from the intestinal tract by Turkish clay and soil which could be a factor

leading to iron deficiency114• They showed that the Turkish clay (20-25 %

montmorillonite and 65-70% sepiolite) had a high CEC and was more effective in

blocking iron absorption than other clay with lower CEC. In their experiments, clay

from New Mexico (99-100% kaolin) had no effects upon iron absorption. In Australia,

although Eastwell (1984)" reported that clay eating could interfere with iron absorption

(section 1.1.6), his study (1979)43 showed no differences in haemoglobin level between

Aboriginal clay eaters and control. The high content of kaolin in the mounds of this

study would support Hausheld's (1975}69 theory that clay c~ating is a traditional practice

more likely to be beneficial to health than dangerous to it (section 1.1.5).

4.2.3 Influence of Age of Mound Material ou Selected Element Concentrations

and Particle Size

In Daly River, the Aboriginal women indicated that they preferred the more recently

built parts of the mounds, they like the taste of it as the old parts are 'not too sweet'

(section 1.1.2.1.1). The reports of possible elemental differences between ages of

termite mound material in the literature are very scarce. Pomeroy (1976Y44 in Uganda

found no significant differences between new and old parts of MacroterlJleS mounds,

even though there was evidence that the old parts had been built several years

previously. His chemical analyses concentrated on organic matter, nitrogen, calcium,

phosphorus, potassium and total exchangeable cations. He reported that the increased

organic matter (from saliva and excretory material) in the mound (compared to the soil}

seems to be chemically stable, as the organic matter content in parts which are several

years old is very similar to that in newly built parts of the mound144• The carbon

180

(organic matter) is positively correlated with the sum of exchangeable cations and acid

extractable phosphorus136, and hence if the organic matter content within the fresh and

old part of the mound remain constant, the sum of cations and acid-extractable

phosphorus may remain similar.

The age effects analyses showed no significant differences in selected element

concentration and particle size for Tumulitermes pastinator and Nasutitermes triodiae

mounds in Daly River site 3 but highly significant differences in aluminium, copper and

iron in Nasutitermes triodiae mounds in Daly River site 4 (Tables 3.7 to 3.9). The

absence of significant differences in selected element concentration and particle size in

site 3 could have been due to the small number of samples and the high standard

deviation from the means, in particular from the calcium mean. The calcium means tend

to be higher in the newly built parts, 32 and 8 % higher inTumulitermes pastinator and

Nasutitermes triodiae mounds respectively. In site 4, although highly significant, the

differences were not major. They were within the standard deviation of the element

mean and were not observed when the statistical analyses were perfonned by positions

(Table 3.12), again probably·because of the lower number of samples. At each position

(top, middle, bottom) in the mound, the general trend was similar with higher

aluminium, copper, iron and potassium in the old parts of the mounds. An increase in

clay could have explained the aluminium, copper and iron increases as they are

positively correlated (section 4.2.2), but there were no significant differences in clay

between ages although the clay means tend to be higher (5 %) in the old part of

Nasutitermes triodiae mounds in Site 4 (Tables 3.11 and 3.12).

It has been reported (section 1.2.1) that the mound is not a static structure, that the

termites continually rebuilt or extend the mound, taking material from the inside of the

mound to the outside. These constant alterations could explain the lack of differences

in elemental composition between new and old parts, with the exception of aluminium,

copper and iron, which each have highly significant lower concentration in the new

material. The reasons why the Aborigines prefer the new parts of the mounds cannot

be related to increases in the major elements or changes in particle size distribution of

new material. It may be simply that the freshly built parts are easier to collect.

181

4.2.4 Influence of Depth in Mound on Selected Element and Particle Size

In accordance with the ways the Aborigines collect the termitaria, a complete study of

the depth effects was not considered necessary, as they collect mainly from the first I 0

em from the outside of the mound. The depth effects was only addressed in the detailed

mound study to have a more complete picture of the mounds studied.

A gradual increase of organic matter and total cations content from outer wall to nursery

area of mounds has been frequently reported, around the world and in Australia, for

many different species27•134

•97

• The extent of the increase (in particular in exchangeable

calcium and magnesium) is closely related to the quantity of organic material

incorporated into the mound and to the type of mound construction. Nevertheless, the

increase in organic matter and total cations have not been observed for all species.

Pomeroy (1976)144, in Uganda, found no difference between the inner and outer parts of

the mound wall of Macrotermes mounds and Okello-Olqya et a! (1985)136 found no

differences in total phosphorus, calcium, JX)tassium, magnesium, sodium and iron in

Amitermes vitiosus and Amitermes /aurensis mounds in Queensland.

The results of the ANOV A for depth effects showed no significant differences m

Amitermes vitiosus mounds (Table 3.5). The results are in agreement with those of

Okello-Oloya eta/ (1985)136• This could be explained by the fact thatAmitermes

vitiosus mounds are homogeneous (section 1.2), whereas in Tumulitermes pastinator

mounds, the inner part of the mound contained significantly higher levels of calcium,

potassium, manganese, clay and silt and lower coarse sand than the outer parts (Table

3.8). The magnesium mean, at depth=2 (0-10 em fraction taken from the inside central

axis of the mound), although not significantly different was 10% higher than at depth=O

(0-1 em fraction of the outside mound) and 13 % higher than at depth= I (0-10 em

fraction of the outside mound). These results are in agreement with the literature values

(Table 1.5). Lee and Wood (197ib)" also noted an increase in clay in the material of

the nursery of Nasutitermes triodiae (Table 1.3). In Nasutitermes triodiae mounds, there

appears to be an increase of calcium at depth=2 (Table 3.10), however the 28% increase

compared to depth= I was not significant because of the high standard deviation from

the calcium mean at depth=2. The high variation of calcium is due to the heterogeneity

182

of the mound composition at that depth. The samples were taken at different levels

within the same depth, one was taken in the nursery (Nt20D3 see Appendix C 1 a and

Clb). The nursery sample had a higher content of calcium (73.6 ± 0.5 mg/lOOg),

magnesium (297.8 ± 2.5 mg/lOOg), manganese (13.5 ± 0.2 mg/lOOg), zinc (2.1 ± 0.01

mg/lOOg), clay (28.1 %) and a lower coarse sand content (15.4 %) than in the other

samples at the same depth. The higher concentration of elements and clay content found

in the nursery is consistent with the data from the literature (see Table 1.3). At

depth=2, the cobalt showed a significant increase but the concentrations were so low that

it is not considered important. The iron decrease at depth=2 was not major although

significantly different, the differences are within the standard deviation of the elements.

The sodium decrease at depth=2 contrasts with the finding of Boyer (1956)" who

reported very high levels of sodium in the inner part of the mound of an African termite

(section 1.2.4.3) and Lee and Wood (1971b)98 findings inNasutitermes triodiae mound

in Daly River where the sodium level was double in the nursery area compared to the

outer galleries (see Table 1.5).

Overall, there are no differences associated with depth in Amitermes vitiosus mounds,

whereas for Tumulitermes pastinator and Nasutitermes triodiae, there are differences

between depths but the differences are principally associated with the different structures

within the mound rather than the depth itself. For example, different structures can be

located within the same depth: nursery and mound galleries can be found in the inner

core of the mound (depth=2).

4.2.5 Influence of Position in Mound on Selected Element Concentrations and Particle Size

In selecting the termitaria sample, the Aborigines did not indicate a preference for any

one position in the mound; however, as part of the overall study, it was decided to

determine whether concentration did in fact vary at different positions in the mound.

The data on variation of element concentration and particle size with mound position are

also very scarce. Okello-Oloya eta/ (1985)136 showed different patterns of variation of

183

selected exchangeable elements and organic carbon between sites according to the

positions in the mound; with the exchangeable calcium levels being higher in the upper

and middle levels of Amitermes Jaurensis mounds. Unlike Okello-Oloyaet a! (1985)136,

Nye (1955)134 reported that the casing of an African termite mound varied little in

composition (organic carbon, pH and exchangeable cations) from the top to the base and

Coventry et a/ (1988)3a found no systematic variation within sampling position in

Amitermes vitiosus, Tumulitermes pastinator and Drepanotermes perniger mounds. In

Daly River, Lee and Wood (1971b)98 found no increase in HCl extractable calcium,

potassium and phosphorus but increase in exchangeable calcium and potassium in the

basal region of Nasutitermes triodiae mound.

Generally in Amitermes vitiosus mounds, ·there seems to be a trend towards higher

concentration of elements in the upper section of the mound (Table 3.6 and 3.14) but

the differences are seldom consistently significant; only calcium, magnesium and zinc

were significantly increased in the top section of mounds_ at two sites: Daly River site

4 and Elliott site 5 (Table 3.14). Highly significant differences were only observed at

Daly River site 4; but those Amitermes vitiosus mounds were not eaten by the

Aborigines. In Tumulitermes pastinator (Tables 3.8 and 3.15) andNasutitermes triodiae

(Tables 3.10 and 3.16) mounds no highly significant differences have been found at all

sites. In contrast to Amitermes vitiosus mounds, calcium tends to accumulate in the

lower section of the mound of Tumulitermes pastinator (Table 3.8) andNasutitermes

triodiae (Table 3.10), but this was not observed at all sites.

No highly significant differences in concentration of selected elements and particle size

composition have been observed according to the position in mounds eaten by the

Aboriginal communities of Elliott and Daly River. Although highly significant

differences were found for Amitermes vitiosus mounds at site 4, these mounds are not

selected for consumption.

184

4.2.6 Influence of Mound Size on Selected Element Concentrations and

Particle Size

The mound size is assumed to be an index of the colony size {although Watson & Perry

(1981)193 have observed that it is not always the case), and larger mounds are assumed

to be older. This is supported by observations of Coventry et a! (1988)38 of a

Tumulitermes pastinator mound which increased in basal diameter from 0.6 to 1.4 m

over a 5 year period and by the fact that very largeNasutitermes triodiae mounds may

be 100 years old. For the purpose of the current study, to compare mounds of different

sizes, it was not necessary to know exactly their volume or age. But as the shape of the

mounds is so variable it was important to take into consideration the fact that some

mounds are tall and narrow and others small and wide. For that reason, the size was

estimated by the sum of the height and the circumference (Figure 3.2). Overall no

significant correlations were found between the size and the selected elements and the

particle size (Table 3.17). This may be linked to the fact that only minor differences

have been observed between old and new materials (section 4.2.3), and very few

differences have been observ~d between positions (section 4.2.5). The termite activities

(constantly re-organising and renewing their habitat) and perhaps as presumed by

Pomeroy (1976)'44 the redistribution of elements throughout the mound by diffusion

during the wet seasons could contribute to the homogeneity of the concentration of

elements and particle size distribution within a mound.

4.2.7 Comparison of Mounds of the same species at the same site

Differences in concentration of elements and particle size between mounds of the same

species at the same site have been poorly documented. In a study by Lee and Wood

(197Ib)518, only one mound per species was taken at each site, with the exception of one

site (Darwin South Port), where twoAmitermes meridiana/is were sampled. In that site,

major composition differences were observed for most of the selected elements. For

example between the two mounds, the potassium varied from 28 to 64 mg/1 OOg, the

phosphorus from 10 to 18 mg/IOOg and the exchangeable magnesium from 9.7 to 14.6

185

mg/lOOg. Important variations also occurred in the particle size, the clay percentage

varied from 13 to 27 %and the fme sand from 42 to 27%. Okello-Oloya eta/ (1985)136

although studying 5 mounds per site, did not discuss mound variation. The high

standard deviations obtained from the means of their mounds for selected element

concentrations could give an indication of the possible variation between mounds

(although other factors could have been involved). The standard deviations from the

means of calcium, phosphorus and magnesium were 68, 44 and 43 % respectively for

Amitermes vitiosus mounds on a site in Queensland. These findings are consistent with

the results of this study indicating at nearly all sites highly significant differences for the

majority of the selected element concentrations for Amitermes vitiosus (Table 3.18). In

Daly River, the Aboriginals seem to select some mounds in preference to others of the

same species at the same site. The reasons could have been based on the differences in

concentration of selected elements between mounds, but it is also equally likely that their

choice could be based on other considerations. For example, in Daly River site 1, they

were not collecting from mounds of Tumulitermes pastinat~r that were attacked by ants

(lridomyrmex sp). However, in our study, 5 of the I5 mounds sampled at site I were

attacked by Jridomyrmex sp but no significant differences were found between mounds

in that particular site.

4.2.8 Comparison Between Different Species Mound Composition at the

Same Site

Although there is an enormous amount of data reporting physico-chemical differences

between species at the same site98•102

, the information is oflimited use in understanding

preferential use by Aboriginals of a particular species over another at the same site. For

example, selection of Tumulitermes pastinator and exclusion of Tumulitermes hastilis

mounds at Daly River site I and selection of Nasutitermes triodiae mounds and

exclusion of Amitermes vitiosus at Daly River site 4. As seen in section I.2.4, those

species are rarely reported and analysed together on the same site.

186

4.2.8.1 Comparison Between Tumulitermes pastinator and Tumulitermes

hastilis Mounds Composition in Daly River Site 1.

There were highly significant differences for the majority of elements studied (Table

3.19). Aluminium, potassium, sodium, zinc and clay were highly significantly increased

and fine sand significantly higher in Tumu/itermes pastinator mounds. Potassium,

sodium, and clay could play an important role in the diarrhoea treatment. As discussed

by Gracey (1991)", the danger from diarrhoea is dehydration which could be caused by

the loss of body fluids and body salts in the diarrhoea fluid. Potassium and sodium

could act as replacement salts and the clay, as reported in section 4.2.2, could have

positive effects alleviating the symptoms of diarrhoea. The highly significant increase

in these particular element concentrations in Tumulitermes pastinator mounds could

contribute to the reason why this particular species mounds are chosen over those of

Tumulitermes hasti/is, by the Aboriginal community of Daly River.

4.2.8.2 Comparison Between Nasutitermes triodiae and Tumulltermes

pastinator MoUnds Composition in Daly River Site 3 and Howard

Springs Site 6.

Although the Daly River Aborigines collected termitaria samples from the two species,

they preferred the Nasutitermes triodiae mounds. At Daly River site 3, there are highly

significant increases in Nasutitermes triodiae, in aluminium. cobalt, iron, potassium,

magnesium, manganese, silt and coarse sand and significant increases in sodium and fine

sand (Table 3.20). Compared to Tumulitermes pastinator mound composition, all the

selected element concentrations mentioned were significantly higher in Nasutitermes

triodiae together with the finer soil fraction (clay + silt) which was 48.8 % in

Nasutitermes triodiae versus 36.8 % in Tumulitermes pastinator. The Daly River

Aboriginal preference for Nasutitermes triodiae mounds could possibly be related to the

higher concentration of the elements or to the increase in the finer texture which could

perhaps make the termitaria more palatable.

187

At site 6 (Howard Springs), the differences between the two species mounds were not

as obvious as in site 3 (Table 3.22). Only calcium and silt were highly significantly

higher in Nasutitermes triodiae than in Tumulitermes pastinator. Compared with site 3

(Table 3.20), the iron concentration in Tumulitermes pastinator was significantly

increased (from 2917 ± 198 to 4527 mg/lOOg) and was, in this site, higher than in

Nasutitermes triodiae mounds, although still within the standard deviation of the means.

There was highly significant increase in silt in Nasutitermes triodiae mounds but unlike

site 3, the finer soil particle (clay+ silt) in the two species mounds have a very similar

ratio: 33.19 and 35.5% in Tumulitermes pastinator and Nasutitermes triodiae mounds

respectively. The major highly significant difference between the two species was with

calcium, where an increase of 67 % in Nasutitermes triodiae mounds was found. The

differences between the two species vary from site to site. These differences can also

be found in the values reported by Lee & Wood ( 1971 b )98 for a few selected elements

in Nasutitermes triodiae and Tumulitermes pastinator mounds in Queensland (Table 1.5).

The values observed were higher in Nasutitermes triodia~ mounds in organic matter,

potassium, clay and coarse sand and the concentrations were superior in Tumulitermes

pastinator in calcium, exchangeables (calcium, potassium, sodium), silt and fine sand98•

In summary, highly significant differences were observed between the two species but

no consistant pattern was noted.

4.2.8.3 Comparison Between Amitermes vitiosus and Nasutitermes triodiae

Mound Composition in Daly River Site 4.

At site 4, the Aboriginals choose Nasutitermes triodiae mounds over Amitermes vitiosus

mounds. Out of the ten selected elements, five were significantly different between the

two species mounds and the increase was always in Amitermes vitiosus mounds. The

only highly significant increases observed in Nasutitermes triodiae mounds were in

particle size fractions where clay and coarse sand were higher in Nasutitermes triodiae

mounds, however, the percentage of the fine fraction (clay+ silt) was similar in both

species (32.9 and 29.9% in Nasutitermes triodiae and Amitermes vitiosus respectively).

188

Only an increase in clay content (22 % in Nasutitermes triodiae versus 16 % in

Amitermes vitiosus) could be related to the Aboriginal preference. Possibly other

considerations could be of importance, sush as the physical aspect of the mound. For

example, the colour of the mounds: dark grey in Amitermes vitiosus and red-ochre in

Nasutitermes triodiae (see Plate 11 & 12) and the solidity of the mounds: concrete like

in Amitermes vitiosus mounds, requiring a stone to chip a part off in contrast to the new

parts of Nasutitermes triodiae mounds which can be easily collected by hand.

4.2.9 Comparison Between Mound Composition of the Same Species at

Different Sites

Highly significant differences between selected elements and particle size have been

observed between sites for each of the three species (Amitermes vitiosus, Tumulitermes

pastinator and Nasutitermes triodiae). For the same species, differences between sites

have been reported in many studies and in particular in Australia for Nasutitermes

triodiae (Table 1.8), Tumulitf.}rmes pastinator (Table 1.9), Tumulitermes hastilis (Table

1.10), Coptotermes acinaciformis (Table 1.6) and Amitermes vitiosus (Table 1.7).

4.2.9.1 Comparison Between Mound Composition of Amitermes vitiosus at

Different Sites

All the selected elements, silt, fine sand and coarse sand were highly significantly

different between the three sites studied (Daly River site 2 and 4 and Elliott). The clay

mean content remained at approximately 16 % at all 3 sites (Table 3.23). In

Queensland, Holt eta/ (1980)78 reported clay mean values of27.5 and 24.4% at 2 sites

and Lee & Wood (197lb)98 in Larrimah (NT) reported a clay mean value of 29 %

(Table 1.3). As different methods of soil fractionation have been used (this study used

the pipette and sieve method described by Coventry and Fett (1979)37 whereas Lee &

Wood (197lb)98 used the method of Hutton (1955) while Holt eta/ (1980)78 did not

189

specify the method (but probably used the sieve and pipette method), it is difficult to

compare the results with each other.

The variations, although highly significant, are not as broad for all the elements studied

(Figure 3.13 and Table 3.23). Aluminium, copper, iron, maguesium, manganese and silt

means varied within a narrow range. Important variations were observed for calcium,

potassium, sodium, zinc and coarse sand, for example, calcium content of 32 ± 9.9

mg/IOOg, 57.2 ± 33.2 mg/!OOg and Ill ± 23.9 mg/IOOg in Daly River site 2, site 4 and

Howard Springs respectively. Okello-Oloya et a/ (1985)136 reported also wide

differences for a number of total elements (Table 1.7). As their method used hot

hydrofluoric acid for extraction it is not possible to compare their results with the results

of this study as a less vigorous type of extraction has been used in this study. A

comparison may be made for iron as a percentage recovery close to I 00 % was

constantly found in this study (section 4.2.1); Okello-Oloya eta/ (1985)136 reported mean

values of 1270 ± 190, 1440 ± 110 and 1510 ± 350 mg/lO_Qg at three sites which are in

agreement with results of this study: 1420 ± 297, !52! ± 198 and 1726 ± 146 mg/IOOg

at sites 2, 4 and 5 respectively.

Differences in selected mineral and particle size composition of Amitermes vitiosus

between sites could possibly indicate reasons of selection/exclusion of a particular site.

For example, Amitermes vitiosus mounds were selected in Daly River site 2 (although

not the first preference) and Elliott site 5 but not in Daly River site 4. However, in

Elliott, no conspicuous mounds of other species could be found around site 5 that could

be have been used instead of Amitermes vitiosus. Between site 4 and site 2, the

observed differences (Table 3.23) would suggest the mounds of site 4 with higher

calcium, potassium, sodium and lower coarse sand content would be more beneficial

than those at site 2. However, there are obviously other factors than the chemical and

particle size content which could influence the choice. For example, site 2 was very

close to the community (4 km away) and in site 4 Nasutitermes triodiae were present

and favoured.

190

4.2.9.2 Comparison Between Mound Composition ofTumulitermes pastinator

and Nasutitermes triodiae at Different Sites

Unlike Amitermes vitiosus, Tumulitermes pastinator and Nasutitermes triodiae have

never been shown to be rejected by the Aboriginals, although highly significant

differences have been observed between both species for all the selected elements and

particle size. The differences between sites in selected element concentrations and

particle sizes for both species were wider than for Amitermes vitiosus (Tables 3.23 to

3.25). For example, the mean iron concentration varied inNasutitermes triodiae from

1343 ± 92 to 2917 ± 198 mg/IOOg in Daly River to 4527 mg/!OOg in Howard Springs.

Between sites, the most obvious difference was for potassium, in both Nasutitermes

triodiae and Tumulitermes pastinator (see Figure 3.13). The differences could have been

due to the difficulty encountered with potassium extraction (low percentage recovery,

section 4.2.1) but the percentage recoveries found forNasutitermes triodiae mound (site

4) and soils (site 4 and 6) compared to the ICP (mixed acid digest with HF) values of

the external laboratory were of the same order (Table 3.2): 64, 60 and 56 %

respectively. The potassium-mean values in Daly River site 4 forNasutitermes triodiae

mounds was 660 ± 68 while in Howard Springs (site 6) it was only 38.5 ± 4.6 mg/IOOg

(Table 3.25).

In Australia, only one set of particle size analyses and two records of chemical analyses

have been reported for Tumu/itermes pastinator (Table 1.3). As the methods used are

so different it is difficult to compare those values with those found in Daly River and

Howard Springs in this present study. As the exchangeable cation method of extraction

is less rigorous than the perchloric/nitric acid extraction, the higher calcium values found

in Queensland (Table 1.9) would indicate a higher calcium content in those mounds.

The clay value (19 %) found in a Queensland mound (Table 1.3) fits within the range

of the values found in Daly River site I (14.8 ± 1.3 %) and Howard Springs (26.9 ± 3.5

%) (Table 3.24), while more variations have been observed between coarse sand values:

4 % in Queensland mounds, but averaging 20.8 ± 0.8 for the three sites (1,3,6) in this

study. The values found in the literature (Table 1.3) must be used with caution as the

method of particle fractionation indicated is different (section 4.2. 9.1).

191

In Australia, values of chemical and physical composition of Nasutitermes triodiae

mounds have been reported by Lee & Wood (1971b)98 (Tables 1.3 and 1.8). They

studied four Nasutitermes triodiae mounds and although their analytical methods are

very different to those used in this study, it is possible to observe wide variations

between sites for a number of elements: organic carbon. potassium, calcium, phosphorus

and exchangeable cations (calcium, potassium, magnesium and sodium). For example

the exchangeable potassium varied from 7.4 to 39.1 mg/lOOg and the exchangeable

sodium from 0.9 to 17.9 mg/IOOg (Table 1.8) at different sites. Major differences were

also observed between particle sizes (Table 1.3). For example, the clay varied from 25

to 36% and the coarse sand from 12 to 30 %. In this study, the clay of Nasutitermes

triodiae mounds varied from 19.4 ± 1.0 to 27.6 ± 3.0% and the coarse sand from 19.2

± 2.5 to 36.9 ± 4.7 %(Table 3.25).

The highly significant differences between particle size found in this study for Amitermes

vitiosus, Tumulitermes pastinator and Nasutitermes triodiae are consistent with the

results of Lee & Wood (1971 b )98 who concluded that there is no evidence that any

individual species had precise requirements of particle sizes for its structures. This is

also supported by the results of Fyfe & Gay (1938)50 for Nasutitermes exitiosus (Hill)

mounds (Table 1.3) in which the clay percentage varied from 11.2 to 37.7 %. This is

in contrast to a number of African tennites such as Apicotermes spp., which use

carefully selected combination of particle sizes in their structures97,

4.2.10 General Overview: Influence of Soil Composition on Composition of Mounds

of Different Species at Different Sites

Nye (1955)134 reported that the chemical composition of tennite mounds seems to vary

according to many factors, amongst them the species of tennite and the site. As seen

in Table 3.26, the selected element concentration and particle size differences between

Amitermes vitiosus, Tumu/itermes pastinator and Nasutitermes triodiae mounds from

different sites are very broad. For example, the minimum ·potassium content in the

mounds studied was 27.7 mg/IOOg and the maximum 956 mg/IOOg. These results are

192

similar to those found by Lee & Wood (197lb)". They reported that the potassium

(from HCl extract) content varied from 21 mg/lOOg in Coptotermes acinaciformis mound

outer casing to 620 mg/lOOg in Coptotermes /actus outer wall (Table 1.5). The

importance of the element concentration and particle size variation between mounds of

the same species or different species could partially be explained by the differences

between the soil (0-10 em) element and particle size contents at the different sites

(Tables 3.26 to 3.28 and Figure 3.14). In Howard Springs, the minimum mean

potassium concentrations amongst the soils studied was 32.0 ± 12.3 mg/1 OOg in Howard

Springs (Table 3.2), amongst Nasutitermes triodiae mounds 38.5 ± 4.6 mg/1 OOg (Table

3.25) and Tumulitermes pastinator mounds 37.3 ± 7.4 mgllOOg (Table 3.24). The same

relationship was found for the maximum mean potassium concentrations, amongst all the

sites studied, the potassium content was the highest in the soil, in Nasutitermes triodiae

mounds and Tumulitermes pastinator mounds at site 3 (Daly River): 694 ± 158, 897 ±

67 and 651 ± 79 mg/lOOg respectively. The same relationship between soil and mounds

was found for the other selected elements (Tables 3.26 and 3.28).

The selected mean element cgncentrations and the fmer soil particle size (clay and silt)

were generally higher (but not always statistically) in the termite mounds studied than

in their adjacent (0-10 em) soils (Table 3.29 and Figures 3.14 and 3.15). At all the sites

studied, the termite mounds had a higher clay content than their surrounding soils

(Figure 3.15 and Table 3.29); the increases were always sigoificantly or highly

significantly different in Tumulitermes pastinator and Nasutitermes triodiae mounds but

no significant differences were found between Amitermes vitiosus mounds and their

adjacent (0-10 em) soil. The majority of studies have shown an increase in clay content

in mounds in comparison with unmodified soils (section 1.2.3.1-B). In Australia, with

the exception of one site where the clay content of the (0-1 0 em) soil horizon was

exceptionally high (53 %), the clay content was always higher in the mounds of the

three species (Amitermes vitiosus, Tumulitermes pastinator and Nasutitermes triodiae)

than in the adjacent (0-10 em) soils (Table 1.3). Lee & Wood (197lb)98 and Holt eta/

(1980)71 reported that the level of clay content in the mounds resembles the content of

deeper horizons which could have levels even higher than in the mounds (Table 1.3).

This finding is important in relation to the medicinal use of termite mounds as the

193

Aborigines prefer termite mounds instead of surface soil. It is easier to take a piece of

termite mound than to have to dig to the subsoil to obtain the same amount of clay,

particularly during the dry season when the soil becomes rock hard. However, during

the wet season, when the termite mound sites are flooded and inaccessible, the Daly

River Aborigines dig the flooded soil close to the mission to collect some subsoil which

has a clay content higher than the monnds (30.2 ± 0.7 %clay, hydrometer method).

According to Nye (1955)134 no general agreement about the differences between the

chemical composition of the mounds and the surrounding soil has been found~ Most

studies throughout the world and in Australia showed an increase in the chemical

composition (organic carbon, nitrogen and exchangeable bases (calcium, magnesium))

of the termite mounds compared to the adjacent soils77•98

•102

• The many factors

responsible for the increase in element concentrations in the mounds have been discussed

in section 1.2.4. However according to a munber of authors (section I .2.4) not all

mounds have a higher element content than their surrounding soils.

The results of this study together with literature show that although the termite mound

composition reflects the composition of the adjacent soil (0~ lOcm), the selected element

content in the monnds was generally higher than in the surronnding top soil (Table

3.29). Nine soil~species pairs out of eleven were significantly higher in aluminium, iron,

magnesium and sodium, eight soil~species pairs were significantly higher in calcium and

seven pairs were significantly higher in potassium and zinc. The increase of element

concentrations in mounds has been attributed to a number of factors (section 1.2.4) such

as the use of richer sub-soil by termites and the incorporation of vegetation, saliva and

excreta (in parts of mounds). The increase could further support the fact that the

Aborigines prefer taking soil from termitaria in preference to the adjacent soil (0~ lOcm).

There are of course other possible reasons for selecting tennitaria, such as, the belief that

soil processed by animals is considered to be safer than that which is not processed100,

194

4.3 Hot Water (''Infusion") Extractable Selected Element Concentrations from

Amitermes vitiosus Mounds (Elliot~ Site 5)

The purpose of the hot water "infusion" extraction was to obtain a measure of the

concentration of selected element present in the "termite mound tea" drunk by the

Aboriginals of Elliott. Potassium, calcium and magnesium were the three principal

elements extracted following hot water "infusion" from mounds with: 10.3 ± 7.83, 6.69

± 3.21 and 2.37 ± 1.55 mg/lOOg respectively (Table 3.30). These concentrations

represent a very small fraction of the nitric/perchloric acid extract: 6.33, 6.37 and 2.56

%. The extraction of these elements was even lower from adjacent (0-10 em) soil

material: 1.84, 1.95 and 0.94 % for potassium, calcium and magnesium respectively.

This higher concentration could indicate that the selected elemental increase in termitaria

"Infusion" extract (Table 3.29) may be of a more available nature as it may come, at

least partially, from tennite by-products (for example, saliva and excreta) which are

more bioavailable. Potassium was the dominant element in the "infusion", and as

mentioned in section 4.2.1, it could be in a different form (more soluble) in Elliott

mounds and soils than at any other sites. The influence of the sample position in the

mound on the selected element concentrations was comparable to the one observed in

the nitric/perchloric extract analyses in regards to the calcium (section 4.2.5 and Table

3.31). A higher calcium concentration was observed in the top section of mounds. A

highly significant increase in iron in the bottom of mounds, after hot water extraction,

may not be very relevant as the iron concentrations were very low.

In comparison to the human recommended dietary intakes (see Table 1.2), the

concentration of selected elements extracted are minimal (Table 3.30) but, nevertheless,

could contribute to the global intakes. For example, if all the calcium present in the

"infusion" was available to the human body, 1.5 to 2.2 litres of "infusion" would be

necessary to cover the daily calcium losses. But in nonnal food, the percentage

absorption of calcium is around 20 %125; therefore, at least five times more 11infusion"

may be needed.

195

Amongst other usages, the "infusion" is given for gastro-intestinal disorders, such as,

diarrhoea. It could help to restore body fluid and a fraction of body salts lost with

diarrhoea fluids and thus prevent dehydration. On the other hand, as the finer fraction

of the soil (clay and silt) is preferentially selected in the infusion, with the heavier

fractions, fine sand and coarse sand, sinking more easily to the bottom and therefore not

drunk, it may possibly be a more selective and pleasant way of eating clay. The heavier

fraction is recycled as poultice and the combination of both ("infusion" and poultice) are

used to bring up milk after birth. In a hot and harsh environment, smearing the poultice

on the chest and back provides a cooling effect due to evaporation and may benefit the

nursing mother, but also, drinking and relaxing while being surrounded by the attention

of family members, could help in releasing oxytocin, (hormone from the pituitary gland

which causes contraction of the muscle fiber of the milk glands, forcing the milk into

larger ducts), therefore promoting the "let-down" reflex. This reflex may be inhibited

if the nursing mother is worried or physically uncomfortable183• The poultice could have

other beneficial aspects as mentioned in the popular literature, to cite a few: clay

poultices placed on the lower abdomen for several days before menstruation prevent

pain; clay increases circulation and flow of oxygen to the skin all over the body; and

miraculous cures have occurred in patients, in a Swiss phthisiology centre, who had their

entire thorax coated with clay41• Unfortunately, no scientific explanations have been

offered for these observations.


Termitaria and Soils Following Pepsin-Hydrochloric Acid Incubation

As discussed in section 1.2, the total amount of element present in the diet is not

necessarily available to the hwnan body. For example, only 5-20% of dietary iron is

absorbed from the diet; and in general, cereal based food products have low iron

availability with absorption of 1-7 %for rice, com and whole wheat flour125• Most

study in Australia has focused on the effects resulting from termite activities in soils

and on the exchangeable or 'available' elements in relation to biological, ecological and

pedological significance of termite modified material. As the primary objective of the

196

analyses of this study was focusing on human nutrition, and no standard bioavailable

methods were established for all the selected elements, a method which would predict

the bio-availability of iron from food was selected. This method, suggested by

Narasinga Rao & Prabhavathi (1978), simulates the digestive conditions in the stomach

(pepsin-HCl, pH !.35) and the intestine (pH 7.5)122• As most elements are mainly

absorbed in the intestine 183, the method was extended to the other elements selected in

this study; this would reflect the potential nutritional value of termitaria more closely as

it would give a more realistic idea of the selected element content present at the

absorption site.

4.4.1 Quality Assurance

The Fe(II}-bipyridine spectrophotometric analysis had a severe interference due to the

high concentration of ferric iron, Fe(III), present in termitaria, it was not possible to

obtain a reliable measure of bioavailable iron, Fe(JI), as the absorbance of the Fe(JI)

a,a.'-bipyridine complex was observed increasing with time. This type of reaction has

previously been described by Lee eta/ (1948) with Fe(II)-1,10-Phenanthroline complex.

They found that the absorbance of the Fe(ll) complex increased with time because the

Fe(Ill) complex is slowly reduced to the Fe(II) complex. They attributed this reduction

to the higher stability constant of the Fe(!!) complex than the Fe(lll) complex". A

similar type of reaction will occur for the Fe(III) and the Fe(II)-u,u' -bipyridine

complexes with time. By complexing the Fe(lll) as [FeF6]3" with potassium fluoride

prior to the addition of the a,a'-bipyridine reagent, it was possible to obtain a reliable

and constant measure of Fe(II), as no increase of the Fe(II) complex was observed.

The effect of pepsin concentration on the selected element concentration was

investigated. A significant increase of aluminium and soluble iron in the pH 7.5 filtrates

for a 0.5 % w/v solution of pepsin, was observed but it was not followed by a

significant increase of the ionisable iron (pH 7.5) (Table 3.32), nor other differences at

all concentrations tested. Therefore, as suggested by Narasinga Rao & Prabhavathi

197

(1978) a 0.5 % w/v solution of pepsin was used in addition to HCl for all the

extractions.

An internal termite mound reference material was used to monitor possible variations of

the selected element composition during analyses. The results showed that the precision

of the method was very high for the selected elements of the pepsin-HCI extracts (Table

3.33) considering the low concentration of certain elements: cobalt, copper and zinc.

The standard deviation from the mean concentration of selected elements (including

soluble iron and ionisable iron) of pH 7.5 filtrates were much higher for a number of

elements: aluminium, copper, iron, manganese and iron (II). It could be due to the

relatively low concentration of those particular elements and to co-precipitation during

neutralisation to pH 7.5.

4.4.2 Selected Element Comparisons ofPepsin-HCl (pH 1.35) Extracts, pH

7.5 Filtrates and Perchloric/Nitric ExtraCts.

A summary of the selected element concentrations of different termitaria and soils in

pepsin-HCl (pH 1.35) extracts, pH 7.5 filtrates and perchloric/nitric acid extracts is given

in Tables 3.35 to 3.40, together with the percentage recovery between treatments. The

graphical comparisons between mounds of different species at the same site and between

sites are given in Appendices Fl-F2.

High element concentrations in perchloric/nitric acid extracts does not necessarily reflect

their bio-availability. A method that simulates the human digestion (pepsin-HCl pH 1.35

followed by neutralisation to pH 7.5) is more likely to represent the quantity really being

extracted in the human stomach and the quantity available for absorption in the intestine.

Pepsin-HCl extractions are far less rigorous than perchloric/nitric extractions and could

result in variations in the ratios of concentrations of element extracted by the two

different methods between different species of termite mounds -at different sites. Some

elements may be in a "more extractable" form in one species than in another or in one

site than in another (see 4.2.1).

198

4.4.2.1 Influence of Age of Mound Material on Selected Element

Concentrations and Particle Size (Depth=O)

As indicated in section 3.4.2.2, the only significant difference between ages of

Nasutitermes triodiae mound materials at site 4, was in soluble iron in pepsin-HCl

extracts (Table 3.38). Although highly siguificant differences had been observed in

aluminium and copper in perchloric/nitric extracts, no differences were observed in

pepsin-HCl extracts and pH 7.5 filtrates. The constant mound modifications by termites

(rebuilding and extending part of the mound, bringing material from inside to the

outside), could explain the lack of differences in concentration of elements between new

and old parts of termitaria.

While in the perchloric/nitric extracts there was a significant decrease in iron in the new

materials, the opposite was observed for soluble iron in the pepsin-HCI extracts where

a significant increase of soluble iron was found in the new material (Table 3.38). No

significant differences in soluble iron and ionisable iron were observed in pH 7.5

filtrates, although their mean~ were higher in the new materials. In the old material the

soluble iron mean was 0.44 ± 0.42 mg/lOOg while in the new material it was 2.24 ± 1.96 mg/IOOg. The lack of significant differences in soluble iron and ionisable iron in

pH 7.5 filtrates was mainly due to the high standard deviations from the means. In

respect to the preference of Aboriginals for the new parts of the mounds over the old

parts, the soluble iron increase in pepsin-HCl extracts in the new parts of termitaria

could be of importance, as it could be related to the Aboriginal usage during pregnancy

where the iron needs are markedly increased (see section 4.4.3). No other increases in

major elements were detected in pepsin-HCI extracts and pH 7.5 filtrates.

4.4.2.2

199

Comparison Between Different Species Mound Composition at the

Same Site

A) Comparison Between Tumulitermes pastinator and Tumulitermes hasti/is

Mounds Composition in Daly River Site 1.

With the exception of aluminium and sodium mean contents in pepsin-HCI extracts,

which were of the same order in both Tumulitermes pastinator and Tumulitermes hastilis

mounds, the mean concentrations of elements were higher in Tumulitermes hastilis than

in Tumu/itermes pastinator mounds (Table 3.35 and Appendix FI). In particular, the

calcium mean was three times higher in pepsin-HCI extracts and pH 7.5 filtrates; the

soluble iron was 16 times higher in Tumulitermes hastilis mounds (0.13 versus 2.07

mg/1 OOg) and the ionisable iron was not detected in the pH 7.5 filtrates of Tumulitermes

pastinator mounds but had a relatively high concentration of 0.54 mg/1 OOg in

Tumulitermes hastilis mounds. The potassiwn which was highly significantly higher in

Tumulitermes pastinator mounds in perchloric/nitric extracts, was 1.5 times higher in

Tumulitermes hastilis mounds in pepsin-HCI extracts and pH 7.5 filtrates.

The concentrations of selected elements in pepsin-HCl extracts and pH 7.5 filtrates

would not appear to be the reasons why the Aboriginals prefer Tumulitermes pastinator

mounds over Tumulitermes hastilis mounds at site I, as the concentrations of elements

were generally higher in the extracts from Tumulitermes hastilis mounds.

B) Comparison Between Nasutitermes triodiae and Tumulitermes pastinator

Mounds Composition in Daly River Site 3 and Howard Springs Site 6.

In the perchloric/nitric extracts (Table 3.20) significant and highly significant increases

were found in Nasutitermes triodiae mounds at site 3 in all but three of the selected

elements (calciwn, copper and zinc). However, only two elements (magnesiwn and

sodiwn) remained higher in pepsin-HCl extracts (magnesiwn and sodiwn) and three in

pH 7.5 filtrates (magnesiwn, soluble iron and ionisable iron) than in Tumulitermes

200

pastinator mounds (Tables 3.37 and Appendix F2). No soluble iron or ionisable iron

was detected in pH 7.5 filtrates of Tumulitermes pastinator mounds and only a small

amount was detected in Nasutitermes triodiae mounds (0.36 ± 0.38 and 0.14 ± 0.17

mg/lOOg respectively). In pepsin-HCl extracts, higher concentrations of copper and zinc

were present in Tumulitermes pastinator mounds and in pH 7.5 filtrates aluminium and

copper contents were higher than in Nasutitermes triodiae mounds. The other selected

element concentrations were of the same order in both species.

The small amount of soluble and ionisable iron found in pH 7.5 filtrates from

Nasutitermes triodiae mounds and the increase in magnesium and sodium could possibly

be related to the Aboriginals preference for Nasutitermes triodiae mounds in favour of

Tumulitermes pastinator mounds for nutritional! medicinal purpose.

In Howard Springs site 6, when comparing Nasutitermes triodiae and Tumulitermes

pastinator selected element composition in perchloric/nitric extracts, it was found that

only calcium was highly significantly higher in Nasutitermes triodiae mounds and

aluminium, iron and manganese were significantly higher in Tumulitermes pastinator

mounds (Table 3.22, Appendix F6). This contrasts with the concentrations of calcium,

soluble iron, ionisable iron, potassium, magnesium and sodium in pepsin-HCI extracts

and calcium. soluble iron, ionisable iron, potassium and magnesium in pH 7.5 filtrates

being higher in Nasutitermes triodiae than in Tumulitermes pastinator mounds (Table

3.40). At site 6, low concentrations of soluble iron (0.03 ± 0.05 mg/lOOg) and no

detectable ionisable iron in pH 7.5 filtrates was found in. Tumulitermes pastinator

mounds.

While no general pattern was found for the concentrations of selected elements between

the two species at the two sites in perchloric/nitric extracts, in the bioavailable extracts,

when differences occurred in element concentrations between the two species, the

majority of increases were found in Nasutitermes triodiae mounds.

201

C) Comparison Between Amitermes vitiosus and Nasutitermes triodiae Mound

Composition in Daly River Site 4.

In the perchloric!nitric extracts the Amitermes vitiosus mounds results indicated higher

element contents (calcium, cobalt, potassium, manganese and sodium: see Table 3.21)

than in Nasutitermes triodiae mounds. In the pepsin-HCI extracts, Amitermes vitiosus

mounds had higher concentrations of aluminium, calcium, cobalt, copper, soluble iron

(1 0 times), ionisable iron (6 times) than the Nasutitermes triodiae mounds while

potassium and sodium were higher in Nasutitermes triodiae mounds (Table 3.39 and

Appendix F4). This contrasts with the concentration of elements between the two

species in pH 7.5 filtrates where out of ten selected elements, seven were found in

similar concentration in both species and three (aluminium, potassium and magnesium)

were higher in Nasutitermes triodiae mounds (Table 3.39).

The Aboriginal preference for Nasutitermes triodiae mounds over Amitermes vitiosus

mounds may possibly be related to the higher potassium and magnesium content in a

potentially bioavailable fonn in Nasutitermes triodiae mounds, to the higher clay content

(section 4.2.8.3) and to the fact that it is physically easier to sample Nasutitermes

triodiae mounds than Amitermes vitiosus mounds.

4.4.2.3 Comparison Between Mound Composition of the Same Species at

Different Sites

A) Comparison Between Mound Composition ofAmitermes vitiosus at Different

Sites

In perchloric/nitric extracts all the selected element concentrations were significantly

different between sites with narrow ranges of variation in aluminium, copper, iron,

magnesium and manganese and larger variations for calcium, potassium, sodium and zinc

(Table 3.29 and section 4.2.9.1). In pepsin-HCI extracts (Tables 3.36 and 3.39;

Appendix F7), a narrow range of variation was observed for aluminium, potassium,

sodium and wide variations were observed for soluble iron (at site 5: 18.2 ± 2.5

202

mg/IOOg and at site 2: !57± 77 mg/IOOg), ionisable iron, calcium (at site 5: 119 ± 15

mg/IOOg and at site 2: 24.2 ± 8.7 mg/IOOg), and magnesium in pH 7.5 filtrates.

In relation to the Aboriginal use (selection of Amitermes vitiosus mounds at Daly River

site 2 and rejection of Amitermes vitiosus mounds at site 4), the differences in selected

element composition do not indicate reasons for selection at one site over another, as the

rejected site (site 4) had higher soluble iron, ionisable iron, calcium and magnesium

concentrations than mounds at site 2.

B) Comparison Between Mound Composition of Tumulitermes pastinator and

Nasutitermes triodiae at Different Sites

In Tumulitermes pastinator (Tables 3.35, 3.37 and 3.40; Appendix F8) and Nasutitermes

triodiae mounds (Tables 3.37, 3.39 and 3.40; Appendix F9), important variations in

selected elements between sites were observed. For example in Tumulitermes pastinator

the soluble iron varied in p~psin-HCI extracts from 8.68 ± 1.43 mg/lOOg in site 6 to

18.0 ± 10.0 mg/IOOg in site 3 (Tables 3.37 and 3.40). In both species, a high content

in perchloric/ nitric extracts was not necessarily followed by a high content in

bioavailable extracts. For example, in the perchloric/nitric extract iron content was

higher in Nasutitermes triodiae at site 6 than at site 4 by a factor of 2.6 and the soluble

iron content was similar in pepsin-HCl extracts and markedly different in pH 7.5 filtrates

where it was 15 times higher in site 4.

4.4.2.4 General Overview: Comparison Between Mound Composition of

Different Species Studied at Different Sites and their Relation to the

Adjacent Soil (0-lOcm)

As in perchloric/nitric extracts (section 4.2.10.2), the majority of selected element

content in the mounds compared to the surrounding soil was higher in pepsin-HCl

extracts and pH 7.5 filtrates (Table 3.43; Appendices Fl-F6). While the differences

between the soil and the termite mound rarely exceeded 150 % in the perchloric/nitric

203

extracts (Table 3.29), it frequently exceeded 200 % in the pepsin-HCl extracts and pH

7.5 filtrates (Table 3.43). In pH 7.5 filtrates, soluble iron was never detected in the

soils but was present in most mounds (Table 3.34). The selected element increases

observed in the mounds (section 4.2.1 0.2, Figure 3.19) have been attributed to different

factors (section 1.2.4) and amongst them the termite use of richer sub-soil material97, and

the incorporation of saliva, excreta and vegetation in the mound by termite activities.

The increased differences between mounds and soils in pepsin-HCI extracts and pH 7.5

filtrates could reflect the higher bio-availability of elements derived from plants and

termite by-products.

Although it is difficult to compare the results of this study with those in the literature

as the methods used are so different (section 1.2.4.3), similar trends have been observed.

For example, Lee and Wood (197lb)98 reported, at Daly River, a higher total potassium

concentration (XRF) in the soil with 2950 mg/lOOg compared to 1730 mgi!OOg in a

Nasutitermes triodiae mound; and the opposite was found following exchangeable

extraction: 5.9 and 39.1 mg/lOOg in soil and mound respectively (Table 1.5). In tltis

study at Daly River site 4 the concentration of potassium in perchloric/nitric extracts was

551 ± 144 mg/IOOg in the soil and 687 ± 44 mg/1 OOg in Nasutitermes triodiae mounds,

which represented a 20 % increase. In the pepsin-HCI extracts, the potassium

concentration was 5.85 mg/lOOg remained in soil and 54.9 mg/lOOg in mounds (or 9

times more in the mounds). In Okello-Oloya et al (1985)136, the same trend was

observed between soil and mounds for sodium but not for potassium where the soil

mound ratio for different analyses (HF and BaCliNH4Cl) remained constant (Table 1.7).

In Davies and Baillie (1988)40 study in Sabah, Northern Borneo, they found that

although aluminium was higher in Macrotermes sp. mounds than soil (515 and 447 ppm

respectively) following digestion in hydrochloric acid after pre-ignition at 800°C. \Vhile

with potassium chloride extraction the levels were much higher in soil than mounds:

33.0 and 3.9 me/lOOg respectively. Similar observations have not been made in this

study, the aluminium in perchloric/nitric extracts was generally higher in mounds than

in soils (Table 3.29); in pepsin-HCl extracts it was higher in the mounds at certain sites

204

and lower in other sites and in pH 7.5 filtrates, when detected, it was higher in mounds

(Table 3.43).

A general increase in 'bioavailable' element concentration in tennitaria compared to the

soils and no soluble iron or ionisable iron being detected in pH 7.5 filtrates from soil

(Tables 3.41 and 3.42) could be contributing factors in explaining the Aboriginal

preference for mounds over soils.

4.4.2.5 'Bioavailable' Composition of Different Mounds at Different Sites in

Relation to Human Needs and Foods.

Vermeer ( 1971) 185 reported very low concentrations of "available" minerals in 0.1 N HCl

extract of a popular Ghanian clay, with calcium, potassium and magnesium values of 12,

16.5 and 3.1 rog/lOOg respectively. Their calcium and magnesium values are close to

those found in this study in soils (Table 3.41) and the potassium value is closer to the

mound value found in this work. In Alabama, Edwards et al (1964)47, analysing some

clay eaten by pregnant women of the rural communities found that 0. 03 mg/1 OOg of iron

and 0.20 mgi!OOg of calcium were available (0.1 N HCl). The quantity of clay

consumed varied from 6 to 130 g per day in Alabama. This is comparable to the

quantity eaten by the Aboriginals in the Northern Territory (section 1.1.2.1.2).

The daily average quantity of termite mound consumption can only provide a small

portion of the RD!s for adults (Table 1.2). For example, if 50 g of termitaria is eaten

(Table 3.41), 9 % of calcium, 7 % of copper and 6 % of magnesium RDis could be

covered. As mentioned in section 1.2 the RDis exceed the actual daily nutrient

requirements to take into consideration the variations in absorption and metabolism. The

daily losses are much smaller, for example, to cover the daily calcium loss in an adult

male, 100-150 mg of calcium are necessary (section 1.2). If all the calcium present in

the pepsin·HCl extracts was available (Table 3.41), one would need to eat 250-350 g of

termitaria daily. To satisfy potassium, sodium and zinc needs, much greater quantities

would be needed.

205

Table 4.1 shows the relationship between termitaria, Aboriginal bushfoods and other

common food is given in Table 4.1. However, one must be very cautious in trying to

compare termite mounds with other Aboriginal bushfoods, not only are the methods of

extraction very different (dry ashing or nitric/sulfuric acid extracts138} but also the

organic composition of food compared to mineral/organic composition of termitaria

render the bushfood more available to humans.

TABLE 4.1 Composition of selected Australian Aboriginal bushfoods30 and Western foods18 in mg per lOOg edible portion.

Bush food Ca Cu Fe K Mg Na Zn

Dioscorea bulbifer 182 0.9 2.8 346 224 287 1.1 (Long yam, raw)

Ipomea graminea 4 0.6 3.0 293 41 31 1.1 (cooked)

Portulaca oleracea 112 0.9 13.0 100 73 20 3.0 (as damper}

Trichosurus 25 0.7 10.3 495 25 147 4.2 arnhemensis (possum, cooked flesh)

Western food Ca Fe K Na

Whole wheat flour 27.6 3.81 312 3.2

Whole milk 120 0.08 160 so Lentils (Boiled) 10.5 2.2 217 9.4

Almonds 247 4.23 856 5.8

Compared to the termitaria minimum, maximum and mean values in pepsin. HCl extracts

(Table 3.41), the calcium, copper and magnesium values in bushfoods are of the same

order, the sodium and potassium are always higher in bushfood and the soluble iron and

ionisable iron are higher in termitaria.

As termitaria are numerous and easily collected, they could act as a supplement for the

elements calcium, copper, magnesium, manganese and iron and complement bushfoods.

206

4.4.2.6 Soluble Iron and lonisable Iron in relation to the human needs

As seen in Table 3.34 and Figure 3.18. only a minute fraction (if any) of the iron

present in perchloric/nitric extract is present in the pH 7.5 filtrates and no significant

differences have been observed between new and old material of termitaria in pH 7.5

filtrates (Table 3.38). In the soil samples. no soluble iron was detected in pH 7.5

filtrates and this was the case for all the sites studied (Table 3.34). This would indicate

that the increased iron found (section 4.2.10.2) in the mounds compared to the soil,

could be of a more bioavailable nature. The iron increase in the mound could originate

from:

a richer sub~soil;

the increased clay, as seen in section 4.2.2, which in the mound is positively

correlated to the clay content;

the organic-rich excreta incorporated in the mound with the excreta coming from

the digestion of plant material.

In contrast to results of Edwards et al (1964t7 results for "available iron" in clay, in

which they indicated that only 0.03 mg/IOOg was available, the results of this study

show markedly higher soluble iron in pH 7.5 filtrates (up to 5.21 mg/IOOg in

Nasutitermes triodiae mound samples at Daly River site 4), (see Table 3.34). However,

what Edwards eta/ (1964t7 called available iron was the quantity of iron at gastric juice

pH (1.35) not at the duodenum pH 7.5. The iron absorption occurs in the duodenum

and upper jejunum where the pH is around 7.51n and therefore a more accurate estimate

of "available iron" would be obtained from the soluble iron present in a pH 7.5 filtrate

as indicated by Narasinga Rao & Prabhavathi (1978)122• If Edwards eta/ (1964)47 were

to have measured the iron at pH 7.5 (by neutralising their pH 1.35 extract), most

probably, no iron would have been detected. As observed (and expected) in this study

as the pH was raised from 1.35 to 7.5 the concentration of iron decreased markedly

(Table 3.34).

207

Narasinga Rao & Prabhavathi (1978)122 calculated the amount ofbioavailable iron using

a "prediction" equation considering the percent ionisable iron at pH 7.5: Y = 0.4827 +

0.4707X, where Y is the percent iron absorption in adult males and X is the percent

ionisable iron at pH 7.5. For example, using data from their research, if the total iron

in rice is 1.8 mg/lOOg, and the percent ionisable iron in pH 7.5 filtrate is 15.0 %, the

percent iron absorption in an adult male is calculated as (0.4827 + 0.4707 x 15) which

equals 7.5 %and the quantity assimilated would be (1.8 x 7.5/100) = 0.135 mg/lOOg;

and a similar calculation for lentils indicates that the quantity of iron assimilated would

be 0.62 mg/1 OOg. As the daily loss of iron for males is 1 mg/day (Table 1.2), these

results would indicate that 740 g of rice or 160 g of lentil would be required to meet the

daily iron needs of adult males.

The Narasinga Rao & Prabhavathi (1978}122 method can be used to ascertain the

bioavailability of elements from edible food which have an in vivo absorption of 1.36

to 3.8 %. Outside that range, the relationship between the in vivo and in vitro methods

needs to be determined. Iron absorption from termitaria does not lie in this range. If

the in vivo iron absorption in termite mound was 1.36 %, this would result for

Tumulitermes pastinator mound sampled at site 1, in 18.5 ± 2.0 mg/lOOg available iron,

or by applying the Narasinga Rao & Prabhavathi (1978)122 "prediction" equation, 38.3

% ionisable iron at pH 7.5; but in this study no ionisable iron was detected in the pH

7.5 filtrates. The termitaria studied had a very high 'total' iron content: 1321-5195

mg/lOOg (Table 3.34) and very low percent of ionisable iron at pH 7.5 (for example,

0.038% for Nasutitermes triodiae mound samples at site 4). If the "prediction" equation

were to be used, it would result in a percent of iron absorbed of 0.4827 + 0.4707 x

0.038 = 0.5 % or 1539 mg/lOOg x 0.5 = 770 mg/lOOg available iron. This clearly

shows that the "prediction" equation is not applicable in this case and the relationship

between the in vivo and in vitro methods still needs to be determined for termitaria.

The quantity of iron available from termite mound might be estimated from the

concentration of ionisable iron present at pH 7.5. For example, it was shown previously

that 0.27 mg/IOOg of ionisable iron from rice gives 0.135 mg/1 OOg of available iron and

1.24 mgllOOg in lentil gave 0.62 mg/IOOg; this indicated that 50% of the ionisable iron

208

was bioavailable. As the form of iron in termitaria is not known, it is difficult to apply

that estimate to clay, although our analyses indicated that the ionisable iron form may

be derived from plant material excreted by the termite and therefore could be similar to

that of other cereal products. On this basis, up to 0.25 mg/1 OOg of iron from termite

mound could be bioavailable to an adult male. This may seem negligible considering

that the Daly River Aborigines eat only around 30-60 g of tennitaria per day. But the

human average daily loss ranges from only 1 mg/day for men and post-menopausal

women to 5-7 mg/day in women during pregnancy. A further consideration is that the

iron bioavailability increases with a number of factors (section 1.2) such as:

-ascorbic acid which is present in many bushfoods30: Terminaliaferdinandiana (Kakadu

plum), Phyllanthus emb/ica (Indian gooseberry) and Dioscorea bulbifera (cheeky yam);

- the iron status of an individual and

- in women during the second and third trimesters of pregnancy.

It is possible, therefore, that even such small amounts could contribute to the overall iron

need of an individual, especially if they are iron deficient and if other dietary input is

inadequate. This may especially be the case with pregnant women, which is when the

termitaria is mostly consumed in Daly River.

CHAPTER FIVE

CONCLUSIONS

209

5 CONCLUSIONS

This study is the first detailed investigation of the possible nutritionaVmedicinal value

of termite mounds consumed by the Aboriginal people. The most common reasons

given by the Northern Territory Aboriginals for termitaria consumption are to treat

gastro-enteric disorders (including diarrhoea) and during pregnancy. This suggests that

a number of elements may be of importance, in particular clay and minerals such as

calcium, iron, magnesium, potassium and sodium. The clay and in particular the kaolin

fraction, may act as an absorbent anti-diarrhoeal and may help to alleviate digestive

disorders. The high concentration of elements in the mounds could provide a potential

nutrient source, mainly during pregnancy when the needs for elements such as iron are

increased.

In respect to the clay content, this study found that at all sites and for all species, the

mounds selected by the Aboriginals had a higher percentage of clay content than the

adjacent top soil (0-10 em). The differences were more pronounced for Tumulitermes

pastinator and Nasutitermes triodiae than for Amitermes vitiosus mounds. The average

clay content of soil was 12.9 ± 4.4 % and for Amitermes vitiosus, Tumulitermes

pastinator and Nasutitermes triodiae mounds was: 16.3 ± 4.6, 20.8 ± 6.2 and 22.8 ±

4.2 % respectively. Interestingly, the species most favoured by the Daly River

Aboriginals (Nasutitermes triodiae), had the highest mean clay content. Differences

occurred between mounds of the same species at most sites and between the same

species at different sites for Nasutitermes triodiae and Tumulitermes pastinator.

However, clay concentration remained at approximately 16% forAmitermes vitiosus at

all sites. In relation to the Aboriginals preference for a particular species at the

exclusion of another at a same site an~ of new material versus old, the study found an

increase in clay content in the selected mound species but no difference was found

between inaterial of different ages.

In the Northern Territory, the dominant mineral in clays is kaolin followed by illites13u.

Mineralogical analyses of termitaria in the Northern Territory have indicated a high

percentage of kaolin type clay with, for example, 65-85 % kaolin in the clay fraction of

210

Nasutitermes triodiae mound in Daly River''. Lee and Wood (197la)" indicated that

the kaolin ratio was higher in subsoil than topsoil. As half of the mounds they studied

and in particular Nasutitennes triodiae mounds came from subsoil, it is not surprising

that the kaolin ratio was higher in mounds than topsoil. The fact that the kaolin appears

to be a major part of the clay fraction of the termitaria studied is supported in this study

by the fact that the clay had a negative correlation with potassium and magnesium 11•

The importance of kaolin in termitaria can be inferred from the fact that it has long been

used for the treatment of gastric-disorders in both traditional and modem

pharmacologies178 (it is the active ingredient in the anti-diarrhoeal medication

kaopectate186). Because of the low cation exchange capacity (less than 10 me/lOOg),

kaolin does not usually interfere with the absorption of iron186•

Undoubtedly, one of the most remarkable aspects of termite mounds is their high

concentrations of elements and in particular calcium, iron, magnesium and potassium.

Unfortunately, only a small proportion of the total element present in termitaria is

capable of being used. For most elements, the bioavailability varies and depends on the

element itself, on the nature of the whole diet and on factors inherent to the individual19•

In this study an in vitro method was chosen to simulate the digestive process of the

stomach (pepsin-HCl digestion, pH 1.35) and that of the small intestine (neutralisation

of the pepsin-HCl extracts to pH 7.5 by NaOH). Generally, the percentage recovery of

calcium between perchloric/nitric extractions and pepsin-HCl extractions from mounds

chosen by Aboriginals was very high (82 %), but for all the other elements it was much

lower with less than 5 % for potassium and less than 2 % for iron. A number of

elements (aluminium, cobalt, copper, iron, zinc) were precipitated during neutralisation

to pH 7. 5. Less than 0.1 % of the iron present in perchloric/nitric extracts is found after

the pepsin-HCl extraction acid neutralisation. Calcium recovery remained high even

after neutralisation (73 %).

Although the termite mound selected element composition reflects primarily the

composition of the adjacent soil (0-lOcm), the selected element content in the

perchloric/nitric extracts was generally higher in the mounds than in the surrounding top

~ --~ -------------

211

soil. The differences between termitaria and soil was even more important in pepsin

HCl extracts and pH 7.5 filtrates. This could reflect the higher bioavailability of termite

byproducts added to the mounds. Overall, in per~hloric/nitric extracts, highly significant

differences in the majority of selected elements were observed between mounds at nearly

all sites but no significant correlations were found between the size of the mounds and

the selected elements concentrations and no highly significant differences in selected

element concentrations were found according to the position in mounds eaten by the

Aboriginal communities of Elliott and Daly River.

The differences in selected element concentrations, in the pepsin-HCI extracts and pH

7.5 filtrates, cannot always explain the Aboriginal preference of a species at the

exclusion of another at the same site or a particular species (Amitermes vitiosus) at one

site (site 2) but not at another (site 4). In relation to the Aboriginal way of sampling

termitaria, that is, preference for new material, no significant differences were observed

between age of mound material; the only exception. being the soluble iron in

Nasutitermes triodiae mounds, which was increased by 44 % in the new parts in the

pepsin-HCl extracts. This contrasted with the perchloric/nitric extract concentrations,

where the iron content was 23 % higher in the old parts. In relation to the Aboriginal

use of termitaria during pregnancy, the increase in soluble iron concentration in the part

of the mound they favoured (new material) could be of importance.

The concentrations of elements from the water "infusion" extract from Amitermes

vitiosus mounds (Elliott, site 5) were minimal compared to the human needs but,

nevertheless, they could contribute to the global intakes, especially calcium. On the other

hand, the finer fraction which includes clay, preferentially selected in the infusion could

be beneficial against gastro-intestinal disorders. The hot "tea" could also provide some

kind of relaxation to the nursing mother and therefore promote the "let-down reflex".

In relation to the human needs, the daily average quantity of termite mound consumption

can only provide a small portion of the RDis for adults. For example, if 50 g of

termitaria were eaten, less than 5 % of the RDis for calcium, copper and magnesium

could be obtained. As the RDis exceed the actual daily nutrient requirements it is

212

possible that a higher fraction of the needs could be covered. It was estimated that up

to 0.25 mg/lOOg of iron from termite mound could be bioavailable to an adult male, as

the daily average loss for men is only 1 mg/day, 30-60 g of termitaria could contribute

to the daily requirements.

There are four factors that have been identified in this study which could contribute to

an explanation as to the Aboriginal preference for termitaria over soils:

a general increase in concentration of selected elements in perchloric/nitric

extracts

an increase in clay content which appeared to be largely composed of kaolin

a higher concentration of 'bioavailable' elements

soluble iron and ionisable iron were present in most mounds whereas not

detected in soils.

There are of course other possible reasons for selecting termitaria in preference to the

adjacent soil (0-1 Ocm), such as,

the belief that soil prOcessed by animals is considered to be safer than that which

is not processed 100;

the inclusion of termite secretions in mounds could also be beneficial: for

example, saliva is used as building material. Unfortunately, little is known of

the saliva composition93 but it sustains larvae, functional reproductives and

soldiers of some species;

the presence of organic constituents: for example, Pomeroy (1983) reported that

denitrifier micro-organisms release nutrients into the mound by their activity;

many fungi are associated with termites, for example, fungi have been found on

the head, abdomen and legs of Nasutitermes sp. m, when the bodies of termites

become contaminated with fungal spores, these spores are often deposited in the

mound158• In Africa, Sands (1970)158 reported that some fungi associated with

termites have been found to possess medicinal properties.

The termitaria eating habit seems therefore to satisfy a number of functions and the

results of the bioavailable analyses indicated that the low concentrations of elements

present could contribute to the RDis, especially if the individual is deficient and if other

213

dietary input is inadequate. But it is also important to remember that there are

deleterious effects in that eating termitaria could be a substitute for food which would

provide more nutrients and cases of partial obstruction of the colon due to excess eating

of termitaria have been reported17•

Since the bioavailability of elements is influenced by a number of factors, a more

complete understanding of the processes will be required involving detailed analyses of

the diet of the Aboriginals, the form of iron present in the termitaria and a study of the

digestive physiology. Tuckerman and Turco (1983)183 indicated that iron(III) is

dissolved in the stomach acid, bound by gastro-ferrin and then reduced to ionisable iron.

The ionisable iron is then absorbed by an active transport mechanism in the duodenum

and upper jejunum and by passive diffusion in the more distal portions of the digestive

tract. As significant concentrations of iron are present in the termitaria (between 7.33

to 245 mg/IOOg of soluble iron in pepsin-HCl extracts) it may be important to determine

how much is bound to gastro-ferrin and reduced to ionisable iron, if this is in fact a

mechanism for iron absorption, as this could provide a significant contribution towards

iron intake.

And finally, as Edwards (1964}47 observed, it would be of interest to explore the possible

presence of reducing substances in clay which may facilitate the absorption of iron from

clay and in this context from termitaria.

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215

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APPENDICES

237

APPENDIX Ia: Particle size of Daly River, site 2, termite mound samples (0-lOcm on the outside of the mound).

Species: Amitermes l'itiorus.

Sample Mound Location Particle size (%of dry weight~

Number@ Number Cia~ Silt Fine sand Coarse sand

Av30D2 I top 17.6 17.7 26.3 41.0

Av31D2 I middle 17.8 17.3 31.2 34.8

Av32D2 I bottom 18.2 19.4 32.4 33.1

Av33D2 2 top 13.1 14.6 35.7 40.4

Av34D2 2 middle 14.3 12.4 37.1 39.9

Av35D2 2 bottom 15.0 12.3 38.4 37.7

Av36D2 3 top 13.7 12.5 26.8 47.2

Av37D2 3 middle 14.9 13.7 29.1 45.3

Av38D2 3 bottom 14.7 15.2 32.3 39.9

Av39D2 4 top 14.2 14.0 25.8 46.5

Av40D2 4 middle 15.6 13.8 34.3 39.0

Av41D2 4 bottom 16.1 14.7 31.6 36.6

@ : Sample number explanation: A• Amirermes vitioJW

3041 Sample number

D2 Daly River, site 2

238

APPENDIX lb: Particle size of Daly River, site 4, termite mound samples (0-lOcm on the outside of the mound).

Species: Amitermes vitiosus.

Sample Mound Location Particle size ~%of dry weight)

Number@ Number Clay Silt Fine sand Coarse sand

Av42D4 I top 8.7 18.2 55.3 17.2 Av43D4 1 bottom !9.2 11.5 42.6 25.6 Av44D4 2 top 13.7 25.8 4&9 15.1 Av45D4 2 bottom 16.5 11.5 43.7 29.6 Av46D4 3 top 11.7 31.3 47.0 9.4 Av47D4 3 bottom !8.9 14.5 36.5 309 Av48D4 4 top 15.2 25.0 41.5 17.1 Av49D4 4 bottom 23.5 12.0 40.7 27.6 Av50D4 5 top 14.6 25.1 46.4 !5.7 Av51D4 5 bottom 18.4 10.1 34.7 36.3 Av52D4 6 top 16.9 22.1 40.8 23.0 Av53D4 6 bottom 21.4 12.0 39.4 27.7 Av54D4 7 top 8.7. 19.8 48.3 22.9 Av55D4 7 bottom 21.5 9.3 44.4 24.0 Av56D4 8 top 21.4 8.4 38.3 27.5 Av57D4 8 bottom 11.7 19.2 44.8 28.4 Av58D4 9 top &6 20.0 44.9 25.8 Av59D4 9 bottom 21.8 8.0 42.3 26.5 Av60D4 10 top 12.4 18.6 39.7 25.5 Av61D4 10 bottom 18.5 11.9 42.7 29.4

@ : Samplenumbere~tplanation: A> : Amiterrrn:s vidosus

2743 : Sample number

04 : Daly River, site 4

239

APPENDIX Ic: Particle size of Elliott (site 5) tennite mound samples (0-lOcm o the outside of the mound unless indicated). Species:

Amitermes vitiosus.

Sample Mound Location Particle size (% of dry weiB_ht~ Number@ Number- Clal: Silt Fine sand Coarse sand

A vOlE I top 24.5 10.6 33.5 35.3

Av02E I bottom 14.9 15.0 33.5 36.5

Av03E 2 top 15.1 13.2 34.0 41.7

Av04E 2 bottom 17.7 10.7 32.2 42.2

Av05E 3 top 13.7 16.0 34.9 37.4

Av06E 3 bottom 9.1 18.7 35.3 39.3

Av07E 4 top 21.8 12.3 32.0 36.2

AvOBE 4 bottom 27.0 4.7 31.8 36.4

Av09E 5 top 6.7 20.1 42.7 36.0

AvlOE 5 bottom 11.7 19.5 35.0 37.8

A vUE 6 top 18.5 11.7 34.4 36.9

Av12E 6 bottom 19.9 11.0 33.8 38.7

Av13E 7 top 12.0 13.2 36.4 41.2

Av14E 7 bottom 14.3 13.6 37.1 40.7

AvlSE 8 top 19.2 16Jl 33.4 35.9 Av16E 8 bottom 17.4 14.5 35.1 35.8

Avl7E 9 top 19.6 11.4 34.1 35.9

AvlSE 9 bottom 18.3 11.7 36.2 36.5

Av19E 10 top 22.5 11.0 32.3 36.4

Av20E 10 bottom 12.2 17.2 39.1 37.3

Av21E 11 top 12.2 17.7 39.1 36.9

Av22E II top 6.0 13.0 47.2 36.5

Av23E II middle 16.0 13.0 31.6 39.3

Av24E 11 middle 22.1 5.5 31.8 39.4

Av25E 11 middle 21.6 7.9 33.4 37.2 Av26E 11 bottom 21.0 5.0 34.7 38.5

Av27E# 11 top 18.3 15.7 33.3 38.5

Av28E# 11 middle 22.6 7.2 33.1 37.4

Av29E# 11 bottom 21.6 6.1 35.1 36.6

@ : Sample number explanation: Av Amitermes vitiosus

01-29 Sample number

E Elliott (site 5)

- : For mound size refer to Table 2.5

• : Sample taken from the middle section or the mound (0-1 Ocm ) •

240

APPENDIX Id: Particle size of Daly River, site 1, termite mound samples (0-lOcm on the outside of the mound).

Species: Tumulitermes pastinator.

Sample Mound Location Particle size {% of dry weiS:ht}

Number@ Number Ctar Silt Fine sand Coarse sand

TpOIDI I top 15.6 8.7 43.4 34.5

Tp02Dl I bottom 13.3 8.7 46.1 33.8

Tp03Dl 2 top 16.7 6.1 38.9 37.7

Tp04Dl 2 bottom 15.0 7.0 38.6 38.9

Tp05DI 3 top 15.1 8.1 39.9 34.6

Tp06DI 3 bottom 16.3 9.8 43.9 28.2

Tp07Dl 4 top 16.4 9.0 428 30.3

Tp08Dl 4 bottom 15.8 9.2 43.4 31.! Tp09Dl 5 top 15.6 7.9 39.9 35.7

Tp10Dl 5 bottom 13.7 9.4 44.0 33.4

TpllDI 6 top 14.0 9.8 44.1 34.8 Tp12Dl 6 bottom 17.5 9.1 43.9 31.8 Tpl3Dl" 6 bottom 14-5 &I 45.9 33.1 Tpi4Dl 7 top 15.6 9.7 424 34.2 Tpi5DI 7 bottom 15.7 9.0 44.7 32.5

Tpi6Dl 8 top 16.2 8.1 38.9 38.4 Tpi7DI 8 bottom 14.4 &3 44.8 33.3 Tpi8Dl 9 top 15.9 9.3 43.8 31.1 Tpi9DI 9 bottom 14.0 7.8 42.7 33.6 Tp20DI 10 top 14.4 7.7 41.1 36.1 Tp21DI 10 bottom 13.1 120 44.0 31.1 Tp22Dl 11 top 12.6 8.9 45.1 31.8 Tp23Dl 12 bottom 14.2 9.1 46.1 28.6 Tp24DI 13 top 13.3 9.7 49.3 27.3 Tpl5DI 14 top 12.6 9.6 47.1 28.7 Tp26Dl 15 top 14.3 8.2 46.2 27.4

@ : Sample number explanation: Tp Tumu.litemaes pa.stinaror

01-26 : Sample number

Dl : Daly River, site 1

• : Newly built mound material

APPENDIX le: Particle size of Daly River, site 3, termite mound samples (O.lOcm on the outside of the mound unless indicated) from one mound of Tumulitermes pastinator.

' Sample Mound Location Particle size {%of d!lweight}

Number@ Number c1ax Silt Fine sand Coarse sand

TJ227D3• 1 top 19.2 16.9 35.5 33.8

TE28D3 1 top 20.1 17.9 37.0 29.7

TE29D3 1 top 18.1 15.9 37.0 33.3

TJ230D3• I top 19.6 16.9 41.3 26.6

TE31D3 I top 19.1 15.7 36.5 29.9

TE32D3 I middle 17.0 16.9 37.3 32.9

TE33D3 1 bottom 17.3 15.5 37.2 31.7

Tp34D3 1 top 22.5 16.9 36.6 27.6

Tp35D3 1 top 19.8 15.3 35.2 30.0

Tp36D3 1 middle 25.0 16.1 37.6 25.4

Tp37D3 1 middle 21.3 18.7 37.5 26.6

Tp38D3 1 bottom 21.0 13.0 34.5 33.5

Tp39D3 I bottom 17.8 13.4 40.3 29.9

Tp40D3# 1 top 20.7 19.1 35.0 27.1

Tp41D3# 1 middle 21.2 19.1 37.4 22.6

Tp42D3# 1 bottom 20.7 19.0 39.7 20.7

Tp43D3## 1 bottom 18.4 17.5 40.4 24.6

@ : Sample number explanation: Tp Tumuliterma pastinator

27-43 Sample number

03 Daly River, site 3

Underlined: Sample collected on the outside of the mound (0-lem)


# : Sample taken from the middle section of the mound (0-lOcm.)

## : Material from the nursery

241

242

APPENDIX If: Particle size of Howard Springs (site 6) termite mound samples (()..10cm on the outside of the mound).


Sample Mound Location Particle size {%of d!l weighQ

Number@ Number Ctar Silt Fine sand Coarse sand

Tp44H 1 top 26.4 6.2 44.5 19.7

Tp45H 1 bottom 23.9 6.4 47.7 20.5

Tp46H 2 top 30.3 5.6 41.9 20.3

Tp47H 2 bottom 29.8 5.5 45.7 16.7

Tp48H 3 top 29.5 6.2 47.1 15.4

Tp49H 3 bottom 27.9 5.0 48.1 16.9

Tp50H 4 top 28.7 6.8 43.7 17.8

Tp51H 4 bottom 23.9 7.6 46.8 19.8

Tp52H 5 top 23.1 6.5 47.4 221

Tp53H 5 bottom 19.7 5.6 48.6 25.3

Tp54H 6 top 28.1 6.6 47.5 17.7

Tp55H 6 bottom 26.2 5.1 49.6 17.3

Tp56H 7 top 23.3 6.1 51.4 16.9

Tp57H 8 top 25.4 5.5 51.0 17.6

Tp58H 9 top 23.4 7.1 49.7 18.9

Tp59H 10 top 25.4 8.0 50.1 17.0

Tp60H 11 top 34.1 8.4 38.3 16.9

Tp61H 11 middle 30.4 7.0 40.7 17.6

Tp62H 11 bottom 31.4 6.7 44.2 16.4

Tp63H 12 top 25.8 5.4 45.8 226

Tp64H 12 middle 23.6 4.1 42.9 26.3

Tp6.5H 12 bottom 24.4 4.4 47.1 21.7

Tp66H 13 top 31.4 4.9 39.0 21.6

Tp67H 13 middle 31.3 6.4 38.8 20.3

Tp68H 13 bottom 24.9 7.6 42.4 23.7

@ : Samplenumberexplanation: Tp Tumu/ilamt:J pasrinator

44-68 Sample number

H Howard Springs (site 6)

243

APPENDIX lg: Particle size of Daly River, site 3, termite mound samples (0-lOcm on the outside of the mound unless indicated) from

one mound of N asutitermes triodiae.

Sample Mound Location Particle size ~%of dry weight}


Nt01D3* 1 top 36.5 12.6 32.7 23.9

Nt02D3 1 top 19.6 29.8 32.4 23.7

Nt03D3* 1 top 18.3 31.0 31.6 20.7

Nt04D3* 1 middle 22.6 28.6 31.4 21.4

Nt05D3 1 middle 18.1 23.8 33.3 20.3

Nt06D3* 1 middle 19.1 28.6 35.8 22.1

Nt07D3 1 middle 22.0 29.1 31.2 20.7

Nt08D3* 1 bottom 21.7 29.2 34.8 19.4

Nt09D3 1 bottom 19.7 19.2 35.4 29.5

Nt10D3 1 top 19.8 284 35.9 19.0

Nt11D3 1 top 18.4 30.4 34.5 18.2

Nt12D3 1 middle 18.7 29.4 33.0 21.4

Nt13D3 1 middle 19.8 .31.6 29.8 21.5

Nt14D3 1 bottom 21.0 28.2 33.7 21.8

Nt15D3 1 bottom 18.7 28.8 34.6 20.7

Nt16D3# 1 top 21.6 29.6 29.5 21.3

Nt17D3# 1 top 19.3 30.0 34.8 20.6

Nt18D3# 1 middle 21.2 31.8 35.0 16.6

Nt19D3# 1 bottom 19.5 32.9 32.7 19.7

Nt20D3## 1 bottom 28.1 24.7 32.7 15.4

@ : Sample number explanation: N< Nasutitermes nWditu

01-20 Sample number

D3 Daly River, site 3

Underlined Sample collected on the outside of the mound (0-lcm)

• Newly built mound material

• Sample taken from the middle section of the mound (0-lOcm)

•• Sample taken from the middle section of the mound (0-lOc:m.) in the nursery

244

APPENDIX Ih: Particle size of Daly River, site 4, termite mound samples (0-lOcm on the outside of the mound unless indicated)

Species: N asutitermes triodiae.

Sample Mound Location Particle size {% of dry weight2

Number@ Number aa~ Silt Fine sand Coarse sand

Nt21D4' I top 22.9 13.2 35.0 34.0

Nt22D4 I middle 24.2 10.3 36.0 31.5

Nt23D4 I bottom 19.7 8.6 36.3 35.8

Nt24D4 2 top 24.8 5.3 30.4 36.9

Nt25D4 2 middle 25.5 7.6 30.1 36.2

Nt26D4 2 bottom 24.2 7.6 34.6 30.9

Nt27D4 2 top 19.0 5.7 31.5 42.6

Nt28D4 2 middle 21.7 8.0 28.0 44.4

Nt29D4 2 bottom 18.0 4.8 29.1 45.8 Nt30D4"' 2 top 21.2 6.8 34.8 35.9 Nt31D4,.. 2 middle 19.6 5.2 34.2 39.1 Nt32D4* 2 bottom 16.1 5.5 36.8 40.1

Nt33D4 3 top 19.5 8.6 31.5 38.5

Nt34D4 3 middle 22.1 6.7 32.3 37.4

Nt35D4 3 bottom 17.3 7.8 36.7 36.7

Nt36D4* 3 bottom 17.8 7.1 39.3 34.6

Nt37D4 3 top 22.3 6.5 36.4 37.3 Nt38D4 3 middle 23.4 5.7 30.2 42.7 Nt39D4 3 bottom 19.5 5.7 31.6 40.9 Nt40D4* 3 top 18.3 6.3 36.4 36.8 Nt41D4* 3 middle 15.7 5.2 38.3 38.3 Nt42D4* 3 bottom 17.3 5.7 33.0 41.3

Nt43D4 4 top 22.3 6.8 29.4 41.4 Nt44D4 4 middle 21.4 5.8 30.8 40.3

Nt45D4 4 bottom 20.9 6.2 33.6 38.1 Nt46D4 4 top ,18.2 5.2 33.7 40.4

Nt47D4 4 middle 16.6 5.2 30.4 46.7 Nt48D4 4 bottom 19.0 5.0 31.6 42.0 Nt49D4* 4 top 18.0 6.8 34.4 43.5 Nt50D4* 4 middle 15.1 6.2 32.3 43.6 Nt51D4* 4 bottom 16.0 6.7 34.5 41.6

Nt52D4 5 top 26.6 7.7 28.3 36.8 continued ...

245

APPENDIX Ih: continued ...

Sample Mound Location Particle size ~%of drywei~hQ


Nt53D4 5 middle 31.1 7.1 28.7 33.1

Nt54D4 5 bottom 22.6 7.0 30.7 37.6

Nt55D4 5 top 24.7 5.2 29.9 36.4

Nt56D4 5 middle 22.6 4.2 30.9 38.5

Nt57D4 5 bottom 22.7 3.8 26.7 42.5

Nt58D4* 5 top 19.7 5.7 31.0 40.8

Nt59D4* 5 middle 19.9 5.8 32.1 39.7

Nt60D4* 5 bottom 19.9 5.6 32.0 41.1

Nt61D4* 6 top 22.8 6.1 30.8 40.7

Nt62D4 6 middle 19.6 7.0 31.0 41.0

Nt63D4 6 bottom 22.8 3.8 29.4 41.2

Nt64D4 6 top 14.0 4.0 37.7 41.6

Nt65D4 6 middle 18.0 6.2 30.6 43.1

Nt66D4 6 bottom 19.6 4.8 28.9 44.6

Nt67D4* 6 top 16.8 6.5 32.0 42.3

Nt68D4* 6 middle 23.9 5.1 25.3 42.7

Nt69D4* 6 bottom 26.8 5.8 27.2 38.7

Nt70D4* 7 top 25.4 4.2 25.3 44.3

Nt71D4 7 middle 27.1 4.8 22.6 44.9

Nt72D4 7 bottom 20.6 6.0 27.3 43.9

Nt73D4 8 top 22.4 7.6 31.3 38.7

Nt74D4 8 middle 17.7 19.8 402 22.3

Nt75D4 8 bottom 22.3 6.7 33.0 37.6

Nt76D4 9 top 19.5 6.5 36.1 40.0

Nt77D4 9 middle 23.8 9.0 33.8 34.4

Nt78D4 9 bottom 21.6 10.0 34.3 35.4

Nt79D4 10 top 22.6 11.6 33.1 35.6

Nt80D4 10 middle 25.5 10.4 34.8 30.4

Nt81D4 10 bottom 25.5 11.7 37.0 25.5

Nt82D4 11 top 25.6 8.4 32.7 34.2

Nt83D4 11 bottom 21.2 7.7 32.1 39.7

Nt84D4 12 top 17.3 6.3 29.7 45.1

Nt85D4* 12 bottom 16.8 8.9 36.7 38.0

continued ...

246

APPENDIX lh: continued ...

Sample Mound Location Particle size (% of d!X weight)

Number@ Number ctar Silt Fine sand Coarse sand

Nt86D4 13 top 19.1 6.3 34.0 39.1

Nt87D4 13 bottom 21.1 6.1 39.4 33.8

Nt88D4 14 top 22.2 8.8 30.5 38.6

Nt89D4 14 bottom 18.6 6.3 36.0 37.7

Nt90D4 15 top 18.8 10.8 37.7 34.8

Nt91D4 15 bottom 20.2 6.9 37.9 34.7

@ : Sample number expla Nt NtmJtiterme:r triodl~

• Newly built mound material


04 Daly River, site 4

Underlined: Sample collected on the outside of the mound (0-lc:m)

247

APPENDIX li: Particle size of Howard Springs (site 6) and Berrimab (site 7) tennite mounds samples (0-lOcm on the outside of the mound).

Species: N asutitermes triodiae.

Sample Mound Location Particle size {%of dryweishq

Number@ Number aa! Silt Fine sand Coarse sand Nt92H I top 23.8 11.0 41.0 25.3

Nt93H* I middle 24.0 7.4 47.3 224

Nt94H I bottom 22.5 6.8 51.8 19.2

Nt95H 2 top 25.4 6.5 48.9 20.8

Nt96H 2 middle 27.0 9.1 45.4 20.2

Nt97H 2 bottom 25.7 8.6 46.8 19.4

Nt98H 3 top 27.5 8.9 43.4 19.8

Nt99H 3 middle 29.0 8.6 45.1 17.2

Nt!OOH 3 bottom 27.6 7.8 44.8 18.6

Nt!OIH 4 top 26.9 7.9 43.6 19.9

Nti02H 4 middle 29.4 7.7 43.2 15.4

Ntl03H 4 bottom 31.6 7.4 43.6 16.5 Nt104H 5 top 30.4 7.3 42.4 18.4

Ntl05H 5 middle 32.7 5.9 42.6 16.1 . Nt106H 5 bottom 30.9 6.9 40.4 19.3 Nt107B I middle 29.1 7.2 44.8 16.1 Nt108B 2 middle 14.9 8.7 44.8 26.2 Ntl09B 3 middle 20.1 11.8 45.9 19.8 Nt110B 4 middle 16.5 10.5 48.0 22.2 Nt111B 5 middle 15.4 11.9 43.2 27.1

@ : Sample number explanation: Nt Nasutitermes trioditu:

92-111 Sample number

H Howard Springs (site 6)

B Berrimah (site 7)


248

APPENDIX Ij: Particle size of termite mound samples (0-lOcm on the outside at the middle height of the mound) at Daly River, site 1:


Sample Mound Particle size ~%of dry weis_ht~

Number@ Number Oay Silt Fine sand Coarse sand

ThOIDI I 8.7 10.2 51.0 29.6

Th02Dl 2 15.2 10.0 46.9 32.3

Th03Dl 3 14.5 9.2 38.3 42.7

Th04Dl 4 11.1 9.2 32.2 49.8

Th05Dl 5 12.3 8.7 26.5 55.6

@ : Sample number explanation: Th ' TJ~mulitermes hastilis


Dl Daly River, site 1

249

APPENDIX Ik: Particle size of soil samples (0-lOcm depth) collected at different sites: Elliott (site 5), Daly River (1-4), Howard Springs (site 6) and Berrlmah (site 7).

Soil Termite Particle size (%of d~ weiB,ht~ Number sEecies@ Clal Silt Fine sand Coarse sand

OlE Av 10.28 5.10 34.61 51.17

02E - 12.03 4.55 29.36 54.09

03E - 12.01 6.85 38.61 43.55

04E - 17.65 4.11 26.34 53.26

05D1 Tp,Th 6.66 9.25 42.18 43.36

06DI - 6.21 9.49 47.32 36.21

07D1 • 6.24 7.11 46.18 40.93

08DI • 6.67 8.03 39.39 43.53

09D2 Av 13.33 13.44 35.02 39.61

10D2 • 15.59 11.44 35.41 40.40

1102 • 13.64 9.24 32.65 43.98

15D3 Tp 10.93 22.06 40.14 30.53

16D3 • 8.19 10.58 39.05 44.45

17D3 Nt 17.44 27.17 40.33 21.58

18D3 • 15.91 26.24 41.29 20.18

19D3 • 17.39 25.75 40.15 20.33

20D3 Tp 7.23 12.08 41.67 40.23

2!D3 Nt 14.1 19.30 44.87 25.02

22D3 Nt,Tp 14.76 18.23 40.09 29.30

23D4 Nt,Av,Ca 14.24 10.34 47.21 32.15

24D4 • 16.35 14.41 47.33 26.21

25D4 • 1241 8.13 39.15 44.51

26D4 • 11.42 8.64 51.65 33.75

27H Nt,Tp 20.59 7.09 46.34 30.73

28H • 22.05 10.52 47.31 24.47

29H • 14.3 6.12 56.12 26.35

30H • 18.93 6.70 54.08 25.72

31B Nt 8.77 11.56 35.83 39.08

32B • 8.3 11.19 58.37 26.83

33B • 13.26 8.90 32.49 39.87

@: Termite species studied at these soil sites

See list of abbreviations p.(xxxii).

Appendix II: Selected elemental composition of termite mounds sampled at Elliott (site 5) following hot water Minfusion~ extraction.

Sample

Number@ A -'liE Av02E A-'l3E A-'l4E A-'l5E Av06E Av07E A-'l8E A,OOE Av10E Av11E Av12E Av13E Av14E Av15E Av16E Av17E Av18E Av19E Av20E Av22E Av26E Av62E Av63E

#OlE (n~l) 02E (n~l)

AI 0.15 .± 0.01 1.34 .± 0.06 0.21 .± 0.20 0.17 .± 0.12 0.07 .± 0.01 1.03 .± 1.45 0.86 .± 0.72 0.53 .± 0.10 0.72 .± 0.11 0.70 .± 0.05 0.56 .± 0.44 0.45 .± 0.43 0.15 .± 0.03 1.22 .± 0.28 0.08 .± 0.11 1.58 .± 1.21 0.97 .± 1.29 0.51 .± 0.24 0.29 .± 0.26 0.29 .± 0.27 0.09 .± 0.13 0.18 .± 0.18 0.20 .± 0.10 0.34 + 0.41

1.18

0.84

Ca

7.26 .± 1.76 2.65 .± 0.45 4.68 .±. 2.51 5.57 .± 0.99 7.72 .±. 2.20 3.99 .±. 1.70 7.83 .±. 0.42 7.51 .±. 2.50 6.09 .±. 2.33 4.60 ± 0.73 8.38 .±. 3.34 1.75 .±. 0.96 6.60 ± 1.62 8.85 .± 1.37

13.52 .±. 7.40 7.10 .± 1.58

13.18 .± 0.20 3.86 .±. 0.83 7.02 .± 2.53 5.34 .± 0.07 4.94 .± 1.16 7.08 .± 1.58

10.73 .± 1.56 9.63 + 2.26

0.97 0.87

Co nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd

Species: Amiter17U!s vitiosus.

Element + sd (n=3) mg1100g

Cu nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd

Fe K 0.03 .± 0.00 22.01 .± 5.49 0.28 .± 0.00 11.49 ± 2.19 0.03 .±. 0.02 0.03 .±. 0.05

nd 0.23 .± 0.17 0.07 .±.'0.10 0.11 .±. 0.03 0.06 .± 0.09 0.23 .±. 0.05 0.09 .±. 0.09 0.12 .± 0.17 0.02 .±. 0.03 0.17 .±. 0.06 0.02 .±. 0.03 0.24 .±. 0.13 0.13 .± 0.19

10.94 .±. 8.78 12.25 .± 4.72 7.01 .±. 5.35 2.15 .± 1.07

14.17 .±. 15.57 3.31 .±. 0.03

13.19 .± 15.51 5.04 .±. 0.05

27.02 .± 3.31 9.11 .±. 6.29 8.16 .±. 1.14 5.06 .±. 3.96 5.05 .± 1.39 3.45 .± 0.39 3.19 .±. 1.10

0.11 .±. 0.01 13.07 .±. 3.61 0.05 .±. 0.07 3.92 .± 1.25 0.13 .±. 0.19 8.64 .±. 5.28 0.03 .±. 0.04 14.39 .±. 2.13 0.04 .± 0.06 16.77 .±. 10.97 0.03 .± 0.04 5.30 .±. 3.73 0.06 + 0.08 4.79 + 2.98

0.12 0.13

1.71 1.85

Mg 3.09 .± 1.00 1.57 .± 0.02 2.23 .±. 1.52 4.58 .± 1.58 3.62 ± 1.47 0. 72 .± 0.46 1.23 .± 0.06 1.39 .±. 0.70 2.45 .±. 1.13 0.65 .± 0.03 3.48 .±. 0.35 1.29 .± 1.09 3.51 .± 0.81 4.06 .±. 4.21 4.21 .±. 2.74 1.16 .±. 0.10 1.92 .±. 0.59 1.95 .±. 0.19 1.70 .±. 1.18 2.17 .± 1.37 1.93 .± 0.48 1.84 .± 0.28 2.06 ± 0.81 2.34 + 1.27

0.47 0.43

Mn nd nd nd nd nd

0.02 .± 0.00 0.01 .± 0.02 0.03 .±. 0.01 0.04 .± 0.05 0.05 .±. o.oo

nd nd nd

0.02 .±. 0.02 0.04 .±. 0.05 0.04 .± 0.03 0.05 .± 0.02

nd 0.04 ± 0.02 0.01 .± 0.01

nd 0.01 .± 0.01 0.07 .± 0.09 0.03 + 0.01

0.02 0.02

@; for explanation of mound sample number refer to APPENDIX Ie.

Detection limit (mg/100g): Co and Zn = 0.02; Cu and Mn = 0.01; Fe = 0.06.

#; for explanation of soil sample number refer to APPENDIX Ik.

Na

1.22 .± 0.95 0.30 .± 0.09 0.20 .±. 0.16 0.27 .± 0.21 0.52 .±. 0.57 017 .± 0.13 1.32 .± 1.51 0.53 .±. 0.32 0.63 .± 0.72 0.18 ± 0.04 1.40 .± 1.11 0.15 .± 0.03 0.29 ± 0.22 0.44 .± 0.32 0.64 .±. 0.50 0.29 .± 0.25 0.13 .± 0.03 0.19 .± 0.03 0.11 .± 0.03 0.27 .±. 0.10 0.22 .± 0.05 0.78 .±. 0.35 0.16 .± 0.04 0.32 + 0.36

0.39 0.36

Zn nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd nd

~ <:>

Appendix Ilia: Selected elemental composition of termite mounds sampled at Daly River, site 2, following perchloric/nitric acid (4:1) extraction. Species: Amilermes vitiosus.

Sample Element + sd {n=3} mll/100g Number@ AI Ca Co Cu Fe K M~ Mn Na Zn Av30D2 3246 .± 91 46.0 .± 0.2 0.70 .± 0.01 0.98 .± 0.02 1773 .± 20 481.2 .± 8.0 126.6 .± 2.6 9.5 .± 0.1 12.6 .± 1.3 1.14 .± 0.05 Av31D2 3205 .± 54 39.8 .± 0.1 0.69 .± 0.01 0.98 .± 0.01 1757 .±4 478.2 .± 31.5 129.2 .± 5.1 8.3 .± 0.0 12.9 .± 1.2 1.15 .± 0.01 Av32D2 3654 .± 100 39.4 .± 0.5 0.73 .± 0.01 1.02 .± 0.01 1856 .± 14 537.9 .± 12.4 138.6 .± 0.7 8.2 .± 0.0 14.5 .± 0.7 0.82 .± 0.71 Av33D2 3034 .±. 94 27.4 .± 0.4 0.48 .±. 0.01 0.69 .±. 0.01 1382 .±. 24 470.4 .±. 443 98.1 .±. 2.1 5.1 .±. 0.1 11.9 .±.OS 0.79 .±. 0.01 Av34D2 3034 .±. 57 21.0 .±. 0.3 0.43 .±. 0.01 0.61 .± 0.01 1255 .± 12 468.7 .± 10.3 90.1 .± 1.4 4.1 .± 0.1 10.8 .± 0.5 0.72 .± 0.02 Av35D2 3091 .±. 47 22.4 .± 1.0 0.42 .±. 0.01 0.60 .± 0.01 1309 .±. 5 486.2 .±. 24.4 95.8 .±. 1.7 4.4 .± 0.1 11.5 .± 0.6 0.74 .± 0.01 Av36D2 2927 .±. 43 50.7 .±. 0.7 0.58 .± 0.00 0.86 .± 0.01 1661 .± 17 410.2 .±. 18.0 107.8 .±. 2.4 10.5 .±. 0.0 12.0 .±. 0.4 1.01 .±. 0.11 Av37D2 3029 .±. 33 28.7 .± 0.2 0.50 .± 0.02 0.74 .± 0.01 1379 .±. 6 426.3 .± 14.9 97.9 .± 1.9 6.6 .±. 0.1 11.3 .±. 0.5 0.83 .±. 0.01 Av38D2 2999 .±. 97 28.6 .± 0.0 0.51 .±. 0.01 0.78 .± 0.01 1514 .± 5 421.4 .±. 27.6 107.2 .±. 2.1 6.8 .±. 0.2 10.8 .± 1.0 0.90 .± 0.03 Av39D2 2%1 .± 77 31.9 .± 0.5 0.53 .±. 0.01 0. 78 .±. 0.01 1478 .± 16 421.4 .± 19.9 98.0 .±. 1.9 7.6 .±. 0.1 11.5 .±. 1.1 0.83 .± 0.00 Av4002 3106 .±. 153 20.7 .±. 0.3 0.49 .±. 0.03 0.74 .±. 0.01 1370 .±. 15 428.8 .± 39.1 98.5 .±. 4.5 5.5 .± 0.1 10.5 .± 0.6 0.82 .±. 0.02 Av41D2 3094 + 53 27.1 + 0.1 0.53 + 0.01 0.81 + 0.01 1517 + 3 429.3 + 23.5 111.6 + 2.3 6.6 + 0.1 10.0 + 0.5 0.92 + 0.01

@;for explanation of sample number refer to APPENDIX Ia.

~ ~

Appendix lllb: Selected elemental composition of termite mounds sampled at Daly River, site 4, following perthlorlc/nitric acid ( 4:1) extraction.


Sample Element + sd (n-3) mg1100g Number@ A1 Ca Co Cu Fe K Mg Mn Na Zn Av42D4 4170 .± 133 97 .± 1.2 0.44 .± 0.01 1.00 .± 0.01 1651 .± 29 782 .± 36 138 .± 2.3 8.6 .± 0.1 33.9 .± 0.8 0.68 .± 0.03 Av43D4 3792 .± 233 36 ± 0.8 0.36 .± 0.02 0.84 .± 0.07 1312 .± 44 702 .± 34 91 .± 4.5 4.5 .± 0.1 27.3 .± 2.3 0.43 .± 0.04 Av44D4 4027 .± 107 92 .± 1.4 0.42 .± 0.01 0.92 .± 0.04 1246 .± 16 680 .± 7 140 .± 1.0 7.3 .± 0.3 28.7 .± 0.5 0.56 ± 0.02 Av45D4 3634 .± 113 24 .± 0.7 0.32 .± 0.01 0.76 .± 0.03 1075 .± 25 649 .± 21 82 .± 4.6 3.3 .± 0.1 25.6 .± 0.8 0.35 .± 0.04 Av46D4 4291 .± 195 102 .± 26 0.47 .± a02 1.20 .± 0.04 2507 .± 84 821 .± 11 152 .± 5.7 9.5 .± 0.2 37.8 .± 3.0 a81 .± 0.04 Av4704 3780 .± 117 35 .± 0.4 0.36 .± 0.02 0.92 .± 0.01 1848 .± 24 682 .± 30 103 .± 1.5 4.7 .± 0.1 26.9 .± 1.5 0.51 .± 0.04 Av48D4 3479 .± 81 100 .± 1.2 0.42 .± 0.01 1.02 .± 0.01 1818 .± 18 728 .± 21 138 .± 2.2 9.1 .± 0.1 34.4 .± 1.3 0.66 .± 0.05 Av49D4 3691 .± 120 48 .± 0.6 0.39 .± 0.01 0.93 .± 0.04 1574 .± 17 732 .± 23 122 .± 2.0 5.3 .± 0.0 28.3 .± 1.4 0.41 .± 0.03 Av50D4 4547 .± 126 133 .± 2.1 0.51 .± 0.01 1.03 .± 0.01 1470 .± 29 911 .± 22 163 .± 1.9 9.2 .± 0.2 42.4 .± 0.6 0.63 .± 0.03 Av5104 3539 .± 34 22 ± 0.5 0.32 .± 0.01 0.66 .± 0.01 1021 .± 5 722 .± 14 84 .± 0.5 3.6 .± 0.0 30.0 .± 0.6 0.40 .± 0.02 Av52D4 3776 .± 55 62 .± 1.0 1.14 .± 0.03 0.88 .± 0.03 1095 .± 10 629 .± 7 119 .± 1.1 5.1 .± 0.1 30.4 .± 1.8 0.48 .± 0.04 Av53D4 3890 .± 181 34 .± 0.2 0.35 .± 0.01 0.74 .± 0.02 1369 .± 3 666 ± 10 100 .± 2.6 4.0 .± 0.0 29.6 .± 0.7 0.38 .± 0.02 Av54D4 3854 .± 52 41 .± 0.9 0.34 .± 0.01 0.85 .± 0.02 1420 .± 31 697 .± 32 112 .± 0.6 4.3 .± 0.1 27.6 ± 1.3 0.41 .± 0.01 Av5504 3788 .±. 230 21 .± 1.2 0.28 .± 0.01 0.75 .± 0.02 1302 .± 27 689 .± 64 92 .± 4.6 2.9 .± 0.1 23.1 .± 1.0 0.36 .± 0.02 Av56D4 3599 .± 88 83 .± 0.3 0.37 .± 0.01 0.86 .± 0.02 1318 .± 18 698 .± 17 126 .± 2.6 6.2 .± 0.0 28.8 .± 1.8 0.40 .± 0.01 Av57D4 4092 .± 56 49 ± 0.7 0.32 .± 0.01 0.78 .± 0.01 1249 ± 31 708 .± 60 111 .± 5.2 4.1 .±. 0.1 33.1 .± 0.2 0.38 .± 0.03 Av58D4 3788 .± 68 52 .± 1.5 0.45 .± 0.02 0.89 .± 0.01 1336 .± 25 694 .± 8 113 .± 3.4 6.4 .± 0.1 33.0 .± 1.1 0.44 .± 0.02 Av59D4 3608 .± 189 31 .± 0.4 0.32 .± 0.01 0.75 ± 0.02 1283 .± 34 690 .± 45 92 .± 6.2 3.4 .± 0.1 31.5 .± 0.3 0.33 .± 0.01 Av60D4 3495 .± 110 64 .± 0.3 0.41 .± 0.00 0.86 .± 0.02 1209 .± 17 701 .± 30 115 .± 5.2 6.8 .± 0.1 29.2 .± 2.6 0.40 .± 0.01 Av6104 3683 + 134 18 + 0.0 0.39 + 0.01 0.74 + 0.02 1064 + 14 702 + 65 88 + 3.8 3.1 + 0.1 34.0 + 0.5 0.40 + 0.01

@: for explanation or ~.ample number refer to APPENDIX lb.

... i!l

Appendix lllc: Selected elemental composition or termite mounds sampled at Elliott (siteS) rollowing perchlodc/nitric acid (4:1) extraction.

Species: Amitermes t>itiosus.

Sample Element + sd (n=3) mg/100g

Number@ AI Ca Co Cu Fe K Mg Mn Na Zn Av01E 4115 .± 117 136.1 .± 1.3 0.36 .± 0.01 0.97 .± 0.00 1790 .± 17 183.1 .± 2.3 99.1 .± 1.0 9.1 .± 0.0 6.6 .± 0.0 1.17 .± 0.09 Av02E 4131 .± 42 132.9 .± 1.3 0.30 .± 0.01 0.96 .± 0.02 1176 .± 9 183.8 .± 0.9 99.7 .± 0.4 8.6 .± 0.1 7.1 .± 0.2 1.12 .± 0.04 Av03E 3557 .± 51 117.9 .± 0.9 0.24 .± 0.01 0.81 .± 0.04 1645 .± 7 154.2 .± 1.9 89.0 .± 0.9 6.0 .± 0.1 6.1 .± 0.1 1.01 .± 0.07 Av04E 3218 .± 69 103.7 .± 0.5 0.27 .± 0.03 0.81 .± 0.03 1588 .± 9 141.8 .± 3.8 81.3 .± 1.0 5.7 .± 0.1 6.0 .± 0.3 1.03 .± 0.03 Av05E 3773 .± 199 87.8 .± 1.0 0.25 .± 0.03 0.91 .± 0.01 1783 .± 9 156.8 .± 1.1 92.3 .± 0.7 4.7 .± 0.1 6.8 .± 0.1 1.02 .± 0.01 Av06E 3585 .t 48 77.5 .± 0.9 0.26 .±. 0.05 0.87 .± 0.01 1726 .± 9 149.3 .± 0.6 89.2 .± 1.2 4.5 .± 0.1 6.6 .± 0.2 0.96 .± 0.02 Av07E 3575 .± 67 104.0 .± 1.5 0.29 .± 0.01 0.91 .±. 0.02 _1832 .t 2 160.9 .±. 5.3 99.1 .± 0.5 6.7 .± 0.2 6.8 .±. 0.3 1.10 .± 0.07 Av08E 3662 .± 8 82.4 .±. 0.8 0.29 .±. 0.02 0.93 .± 0.01 1866 .± 5 168.2 .± 3.9 97.0 .± 0.6 6.5 .± 0.0 6.4 .± 0.1 1.10 .± 0.01 Av09E 3972 .± 20 93.4 .± 0.7 0.40 .± 0.01 1.05 .± 0.01 1909 .± 9 175.3 .± 1.4 99.5 .± 0.1 11.5 .± 0.1 6.3 .± 0.2 1.20 .± 0.00 Av10E 3610 .± 198 76.4 .± 1.4 0.37 .± 0.01 0.95 .± 0.02 1769 .± 8 161.3 .± 2.5 87.1 .± 23 10.2 .± 0.2 5.7 .± 0.0 1.13 .± 0.03 AvllE 3792 .±. 52 119.3 .± 0.5 0.31 .± 0.01 0.96 .± 0.01 1896 .± 18 163.3 .± 0.8 96.3 .± 0.7 8.7 .± 0.1 7.7 .± 0.5 1.23 .± 0.19 Av12E 3805 .± 74 95.5 .± 0.6 0.31 .± 0.03 0.92 .± 0.00 1878 .± 6 164.2 .± 3.7 94.0 .± 0.6 7.7 .± 0.1 7.5 .± 0.3 1.11 .± 0.02 Av13E 3543 .± 11 114.4 .± 1.1 0.27 .± 0.02 0.81 .± 0.01 1611 .± 5 153.1 .± 6.0 91.8 .± 0.4 5.8 .± 0.1 6.3 .± 0.1 0.99 .± 0.04 Av14E 2954 .± 93 91.9 .± 0.7 0.20 .± 0.02 0.74 .± 0.02 1477 .± 6 142.7 .± 25 80.2 .± 1.0 5.7 .± 0.2 5.7 .± 0.6 0.92 .± 0.03 Av15E 4770 .± 118 109.9 .± 1.5 0.31 .± 0.00 1.03 .± 0.01 2127 .± 14 186.6 .± 2.6 113.5 .± 0.4 5.9 .± 0.0 8.4 .± 0.3 1.17 .±. 0.03 Av16E 3942 .± 187 72.8 .± 1.2 0.27 .± 0.03 0.89 .± 0.01 1846 .± 15 166.6 .± 7.4 91.3 .± 2.1 4.8 .± 0.1 7.0 .± 0.1 1.04 .± 0.03 Av17E 3334 .± 53 139.4 .± 2.4 0.34 .± 0.00 0.95 .± 0.01 1589 .± 31 164.8 .± 2.8 91.8 .± 0.7 10.1 .± 0.2 5.5 .± 0.3 1.14 .± 0.02 Av18E 3173 + 80 126.1 + 2.1 0.33 + 0.02 0.89 + 0.01 1504 + 15 154.2 + 1.5 87.3 + 1.5 8.9 + 0.1 6.0 + 0.3 1.06 + 0.04 - -- - ------Av19E 3913 .± 88 105.9 .± 1.4 0.35 .± 0.02 1.01 .± 0.00 1765 .± 8 170.9 .± 1.0 96.0 .± 1.7 &5 .± 0.0 6.2 .± 0.2 1.15 .± 0.01 Av20E 3494 .± 109 86.0 .± 1.9 0.35 .± 0.01 0.93 .± 0.01 1635 .± 34 155.7 .± 3.2 82.7 .± 1.3 9.0 .± 0.1 6.3 .± 0.2 1.08 .± 0.02 Av21E 3425 .± 8 135.4 .± 1.0 0.36 .± 0.01 0.91 .± 0.01 1702 .± 22 155.2 .± 1.2 91.8 .± 0.1 8.3 .± 0.2 6.5 .± 0.4 1.12 .± 0.05 Av22E 3479 .± 129 135.4 .± 4.6 0.30 .± 0.00 0.88 .± 0.02 1575 .± 39 155.9 .± 5.8 93.0 .± 2.9 7.8 .± 0.3 6.1 .± 0.2 1.15 .± 0.06 Av23E 3396 .± 106 154.6 .± 4.1 0.29 .± 0.00 0.94 .± 0.03 1628 .± 15 152.8 .± 2.1 92.2 .± 3.3 8.0 .± 0.1 6.3 .± 0.4 1.11 .± 0.01 Av24E 3607 .± 11 142.3 .± 1.2 0.31 .± 0.01 0.97 .± 0.06 1672 .± 47 148.9 .± 9.2 93.5 .± 0.6 7.5 .± 0.1 6.5 .± 0.1 1.16 .± 0.02 Av25E 3448 + 24 143.9 + 0.4 0.34 + 0.02 0.90 + 0.01 1643 + 32 155.1 + 2.8 93.1 + 1.1 8.3 + 0.1 6.0 + 0.1 1.12 + 0.01 - -- - ------Av26E 3213 .± 69 104.1 .± 0.5 0.31 .± 0.02 0.89 .± 0.01 1642 .± 21 156.8 .± 3.3 87.8 .± 1.9 _7.6 .± 0.1 5.7 .± 0.2 0.90 .± 0.04 Av27E# 3682 .± 28 129.4 .± 1.2 0.35 .± 0.02 0.91 .± 0.02 1688 .± 20 161.5 .± 4.8 96.4 .± 1.0 7.5 .± 0.2 6.5 .± 0.4 1.18 .± 0.05 Av28E# 3634 + 81 147.9 + 2.2 0.34 + 0.01 0.91 + 0.01 1668 + 6 156.8 + 2.3 94.0 + 1.5 7.9 + 0.0 5.8 + 0.2 1.15 + 0.02 - -- - ------Av29E# 3329 + 88 105.0 + 3.8 0.36 + 0.02 0.96 + 0.02 1641 + 34 162.3 + 4.7 86.0 + 2.6 7.8 + 0.2 5.7 + 0.1 0.98 + 0.03

@: for explanation of sample number refer to APPENDIX Ic

.... 11!

Appendix Hid: Selected elemental composition of termite mounds sampled at Daly River, site 1, following perchloric/nitric acid (4:1) extraction.


Sample Element + sd (n=3) mg/100g Number@ AI Ca Co Cu Fe K Mg Mn Na Zn Tp01D1 3306 .± 200 31.3 .± 0.9 0.28 .± 0.01 0.60 .± 0.01 1597 .± 40 549 .±50 85.9 .± 4.4 4.0 .± 0.2 33.4 .± 0.4 0.46 .± 0.02 Tp02D1 2676 .±. 108 23.0 .±. 0.3 0.28 .±. 0.00 0.53 .± 0.01 1248 .±. 16 476 .±. 6 58.3 .±. 2.2 3.4 .± 0.1 24.6 .±. 1.7 0.46 .± 0.02 Tp03D1 2885 .± 73 25.6 .± 0.3 0.29 .± 0.00 0.55 .± 0.01 1228 .± 29 526 .± 24 64.9 .± 0.9 3.1 .±. 0.0 24.0 .±. 1.6 0.49 .± 0.03 Tp04D1 2882 .± 59 25.7 .±. 0.4 0.28 .± 0.01 0.54 .±. 0.01 1201 .± 3 500 .±. 13 64.3 .±. 0.9 3.3 .±. 0.0 24.4 .± 1.1 0.53 .±. 0.03 Tp05D1 3257 .± 130 15.2 .±. 0. 7 0.29 .± 0.00 0.55 .± 0.02 1331 .± 51 490 .± 25 65.3 .± 22 28 .± 0.0 29.3 .± 1.3 0.53 .± 0.02 Tp06D1 3257 .± 83 20.3 .± 0.2 0.28 .± 0.01 0.56 .±. 0.02 1345 .± 20 530 .± 27 67.3 .± 0.6 3.3 .± 0.0 28.7 .± 0.8 0.49 .± 0.02 Tp07D1 3408 .± 84 23.9 .±. 0.3 0.31 .± 0.01 0.63 .±. 0.02 1389 .± 19 557 .± 16 69.8 .±. 2.0 3.1 .±. 0.0 30.2 .± 1.6 0.50 .± 0.03 Tp08D1 3069 .± 70 26.1 .± 0.2 0.28 .±. 0.02 0.58 .±. 0.01 1313 .±. 8 504 .± 14 67.8 .±. 0.5 3.1 .± 0.0 26.8 .± 1.5 0.47 .± 0.02 Tp09D1 2891 .± 12 22.3 .± 0.2 0.28 .± 0.01 0.55 .±. 0.01 1260 .± 11 502 .± 12 61.1 .± 0.1 2.9 .±. 0.1 26.1 .± 0.8 0.46 .±. 0.01 Tp10D1 2849 .± 38 26.8 .±. 0.3 0.27 .±. 0.01 0.56 .± 0.02 1245 .±. 12 497 .± 21 64.5 .± 1.1 3.3 .± 0.0 26.2 ·± 0.6 0.49 .±. 0.05 Tp11D1 2911 .± 103 22.2 .± 0.2 0.27 .± 0.01 0.55 .± 0.00 1337 .± 11 502 .±. 27 61.9 .± 1.3 3.1 .±. 0.1 27.3 .± 2.4 0.48 .± 0.01 Tpl2D1 3270 .±. 69 48.8 .± 0.5 0.32 .± 0.01 0.62 .±. 0.01 1476 .± 9 511 .± 27 83.2 .±. 1.6 4.6 .±. 0.0 27.6 .±. 1.1 0.51 .± 0.01 Tp13D1* 2676 .± 125 26.1 .±. 0.2 0.27 .±. 0.01 0.57 .± 0.01 1329 .± 24 4% .± 4 64.9 .± 1.9 3.5 .± 0.0 25.8 .± 23 0.45 .± 0.02 Tp14D1 3353 .± 11 22.2 .± 0.3 0.29 .± 0.02 0.65 .± 0.01 1422 .± 9 528 .±. 8 68.9 .±. 0.6 3.0 .±. 0.0 28.8 .± 0.2 0.45 .± 0.02 Tp15D1 2949 .±. 146 25.1 .±. 0.9 0.28 .±. 0.01 0.59 .±. 0.03 1373 .± 26 518 .± 22 66.5 .± 2.1 3.2 .± 0.1 27.1 .± 3.2 0.45 .± 0.02 Tp16D1 3066 .± 104 15.2 .±. 0.1 0.31 .± 0.02 0.58 .± 0.00 1533 .± 12 476 .± 23 61.7 .± 21 3.3 .± 0.0 24.5 .± 3.4 0.49 .± 0.01 Tp17D1 2880 .± 10 25.7 .± 0.6 0.28 .± 0.01 0.55 .± 0.01 1398 .± 36 536 .± 23 64.2 .± 0.8 4.1 .±. 0.0 25.4 .± 1.5 0.50 .± 0.02 Tp18D1 3063 .± 123 24.4 .±. 0.2 0.30 .± 0.00 0.56 .±. 0.01 1365 .± 29 507 .± 16 65.2 .± 0.6 3.7 .± 0.1 25.4 .±. 2.0 0.56 .± 0.04 Tp19D1 2892 .± 20 19.3 .±. 0.1 0.30 .± 0.02 0.52 .± 0.01 1311 .± 7 486 .± 37 60.8 .± 0.4 3.8 .± 0.0 25.0 .± 0.2 0.50 .± 0.02 Tp20D1 2875 .± 140 26.7 .± 0.2 0.28 .± 0.01 0.58 .± 0.01 1284 .± 22 492 .± 38 57.3 .± 1.7 3.1 .± 0.1 27.3 .± 4.6 0.48 .± 0.02 Tp21D1 2958 ± 64 27.2 .± 0.3 0.29 ± 0.00 0.59 .±. 0.01 1287 .± 8 481 ± 10 60.3 .± 0.8 3.6 ± 0.0 29.0 ± 2.9 0.48 .± 0.03 Tp22D1 3142 .± 52 17.1 .±. 1.2 0.29 ± 0.02 0.55 ± 0.01 1380 .± 9 520 .± 18 57.8 .± 1.0 3.2 .± 0.1 27.7 .± 1.0 0.50 .± 0.01 Tp23D1 3137 ± 78 26.3 .± 0.4 0.29 .±. 0.02 0.57 .± 0.01 1306 .± 20 517 .±. 27 69.4 .± 1.9 3.1 .± 0.1 31.2 .± 1.7 0.42 ± 0.03 Tp24D1 3063 .± 52 16.4 .± 0.1 0.29 .± 0.02 0.54 .± 0.00 1376 .± 11 505 .± 24 57.8 .±. 0.9 3.1 .±. 0.1 27.9 .± 0.9 0.49 .± 0.01 Tp25D1 3165 .±. 100 13.0 .± 0.0 0.28 .± 0.01 0.52 ± 0.01 1249 .± 4 504 .± 15 58.9 .± 2.0 2.9 .± 0.1 27.3 .±. 3.0 0.46 .± 0.02 Tp26D1 3254 .± 11!__19.7 + 0.3 0.~.01 0.57 + 0.01 1327 + to 544 __..±.___] 58.2 + 1.3 2.8__.±. 0.1 27.0 .±. 3.0 0.49 .± 0.01

@:for explanation of aample number refer to APPENDIX Id.

"' :c

Appendix llle: Selected elemental composition of tennite mounds sampled at Daly River, site 3, following perchloric/nitrlc acid (4:1) extraction.

Species: Tumuliternres pastinalor.

Sample Element ± sd (n=3) mg/100g Number@ AI Ca Co Cu Fe K Mg Mn Na Zn Tp27D3• 3873 .±. 102 18.7 .±. 0.2 0.44 .±. 0.01 0.76 ± 0.02 2835 .±. 20 708 .± 37 148.1 .± 4.7 5.3 .± 0.1 14.2 .± 1.1 1.16 .± 0.02 Tp28D3 4414 .± 292 16.7 .± 0.4 0.46 .± 0.01 0.81 .±. 0.02 3253 .± 46 746 .± 1 156.7 .± 7.6 5.3 .±. 0.1 15.4 .± 0.2 1.13 .± 0.04 Tp29D3 3790 .±. 147 17.8 .±. 0.3 0.43 .± 0.02 0.96 .± 0.04 2859 .± 19 712 .±. 33 148.1 .±. 4.4 5.2 .± 0.1_ 14.5 .± 1.5 1.25 .± 0.04 Tp30D3• 4100 .± 146 29.5 .± 1.2 0.46 .± 0.00 0.94 .± 0.01 2988 ± 68 803 .± 32 181.4 .±. 6.2 6.6 .±. 0.2 16.4 .± 1.1 1.35 .± 0.03 Tp31D3 4607 .±. 51 15.5 ± 0.2 0.50 .±. 0.04 0.87 .± 0.02 3159 .± 4 811 .± 20 171.1 .± 2.4 5.9 .± 0.1 19.8 .± 0.6 1.27 .± 0.03 Tp32D3 3596 .± 55 14.9 .± 0.4 0.42 .±. 0.01 0.76 .± 0.01 2744 .± 32 619 .± 8 146.1 .± 0.4 4.9 .±. 0.1 12.7 .± 0.4 1.15 .± 0.04 Tp33D3 3506 .± 165 17.7 .± 0.2 0.39 .± 0.01 0.87 .± 0.01 2696 .±. 28 617 .± 43 152.8 .± 6.0 5.8 .±. 0.1 12.1 .±. 1.2 1.37 .±. 0.04 Tp34D3 3630 .± 183 24.6 .± 0.4 0.39 .± 0.02 1.67 ± 0.05 2677 .± 37 593 .± 40 162.4 .± 6.8 5.5 .± 0.1 10.6 .±. 1.5 1.66 .±. 0.05 Tp35D3 3959 .± 263 22.9 .± 0.4 0.45 .± 0.02 0.76 .± 0.04 2965 .± 75 664 .t 3 166.7 .± 10.1 6.6 .± 0.2 15.6 .±. 1.1 1.26 .±. 0.02 Tp36D3 4010 .± 157 18.6 .± 0.3 0.40 .± 0.01 1.64 .± 0.02 2993 .± 87, 670 .±. 23 144.6 .± 4.9 5.0 .± 0.1 13.9 .± 0.3 1.73 .±. 0.01 Tp37D3 4550 .± 2 21.0 .± 0.4 0.48 .± 0.03 1.99 .± 0.08 3242 .± 68 794 .± 22 174.0 .± 0.7 5.6 .± 0.1 16.5 .± 0.6 1.76 .± 0.04 Tp38D3 3506 .± 196 22.4 .± 0.7 0.39 .± 0.01 0.90 .± 0.01 2775 .± 6 607 ± 40 127.6 .± 4.0 5.7 .± 0.0 '12.7 .± 1.4 1.13 .± 0.03 Tp39D3 3301 .±. 160 30.0 .± 1.8 0.39 .± 0.02 2.06 .± 0.13 2852 .±. 104 579 .± 63 135.6 .± 7.0 7.3 .± 0.2 13.5 .±. 1.1 1.72 .± 0.07 Tp40D3# 3775 .± 35 45.4 .± 0.6 0.49 .±. 0.01 1.45 .±. 0.04 3023 .± 35 805 .± 27 160.9 .± 1.2 8.4 .±. 0.0 17.2 ± 0.5 1.68 .± 0.01 Tp41D3# 4403 .± 73 56.0 .±. 3.7 0.47 .± 0.01 1.84 .± 0.02 3040 .±. 18 855 .±. 56 186.1 .± 2.3 10.2 .± 0.0 18.4 .± 0.7 1.94 .± 0.05 Tp42D3# 4058 .± 55 84.6 .± 0.2 0.52 .± 0.02 1.36 .± 0.05 3031 .± 43 759 .± 31 192.7 .±. 2.0 12.8 .± 0.1 16.3 .± 1.1 1.81 .± 0.01 Tp43D3## 3919 .± 169 53.9 ,± 0.5 0.43 .± 0.03 0.73 .± 0.01 2660 .± 31 714 + 30 161.6 + 6.5 8.6 + 0.2 15.0 .± 1.1 1.22 .± 0.01

@: for explanation of sample number refer to APPENDIX I e.

.... i::

Appendix III£: Selected elemental composition of termite mounds sampled at Howard Springs (site 6) following perchloric/nitric acid (4:1) extraction. Species: Tumulitermes pastinator.

Sample Element + sd (n=3) mg/100g Number@ AI Ca Co Cu Fe K

Tp44H 6230 ± 29 33.4 ± 0.9 1.08 ± 0.02 2.43 ± 0.01 4894 ± 75 53.4 ± 0.4 Tp45H 5982 ± 49 27.2 .± 0.8 1.07 ± 0.02 2.40 .± 0.03 5800 ± 35 50.8 ± 1.1 Tp46H 7088 .± 14 37.8 ± 0.3 1.20 .±. 0.01 2.02 .±. 0.02 4882 .±. 29 39.1 .± 1.3 Tp47H 6873 .± 55 38.6 .± 0.1 1.17 .± 0.00 1.97 .± 0.01 4128 .± 13 37.5 .± 0.7 Tp48H 6284 .±. 31 55.8 ± 0.7 1.10 ± 0.01 1.91 .±. 0.08 3746 ± 25 36.1 .±. 0.6 Tp49H 6062 .±. 29 48.3 ± 0.4 1.09 .±. 0.01 1.81 .±. 0.01 3713 ± 28 34.7 .±. 0.5 Tp50H 5989 .±. 47 70.1 .±. 0.6 0.73 .± 0.01 1.53 .±. 0.01 4303 .±. 22 34.4 .± 1.9

"Tp51H 5717 .±. 90 62.4 .± 1.0 0.70 .± 0.02 1.46 .± 0.04 4795 .± 85 34.6 ± 0.5

Mg Mn 47.5 ± 0.8

"45.9 .±. 0.4 55.4 .± 1.4 56.2 ± 0.5 50.3 .± 1.1 51.2 .± 1.1 44.9 .±. 1.9 47.0 + 0.6

6.4 .±. 0.0 7.7 .±. 0.1 5.3 .± 0.0 4.9 .±. 0.0 5.5 .± 0.1 5.5 .±. 0.1 5.9 .± 0.1 6.9 .±. 0.2

Tp52H 5684 .±. 63 48.2 .±. 0.5 0.78 .±. 0.02 1.46 .±. 0.01 5399 ± 57 27.7 .±. 0.5 42.7 .±. 0.1 8.7 .±. 0.1 Tp53H 5228 .± 51 54.1 .± 0.5 0.71 .± 0.01 1.38 .± 0.01 6519 .± 119 27.7 .± 0.6 41.7 .± 0.1 10.9 .± 0.1 Tp54H 6639 .±. 137 52.3 .±. 1.2 1.11 .± 0.03 1.84 .± 0.02 4581 .±. 48 37.1 .± 0.7 50.2 .±. 0.3 6.2 .± 0.0 Tp55H 6335 .±. 160 47.0 .± 0.5 1.07 .± 0.03 1.78 .± 0.02 4887 .± 61 35.0 .± 1.0 50.8 .± 1.5 6.9 .±. 0.2 Tp56H 5654 .±. 74 46.7 ± 0.6 1.21 .±. 0.01 1.85 .±. 0.01 2895 .±. 34 48.4 .±. 0.8 50.5 .±. 1.7 6.7 .± 0.0 Tp57H 5644 .±. 123 73.2 .±. 2.4 1.02 .±. 0.03 1.60 .±. 0.01 3934 .±. 45 38.9 .± 0.7 47.0 .±. 0.8 7.2 ± 0.1 Tp58H 5773 .±. 70 51.1 .±. 1.8 1.01 .± 0.01 1.49 .± 0.01 4306 .± 12 29.3 .± 1.3 46.5 .±. 1.1 11.0 .±. 0.0

Na 6.3 .±. 0.3 6.9 .± 0.2 6.7 .±. 0.4

Zn 1.38 .±. 0.01 1.36 .± 0.03 1.31 .± 0.04

6.5 .± 0.3 1.24 .± 0.02 6.8 .± 0.1 6.6 .± 0.2 6.1 .±. 0.3 6.9 .±. 0.0 5.6 .± 0.2 5.6 .± 0.1 6.2 .± 0.6 6.6 .± 0.4 6.2 .±. 0.0 5.6 .±. 0.4 6.3 .± 0.3

1.37 .±. 0.03 1.62 ± 0.17 1.10 .± 0.01 1.05 .± 0.04 1.00 .± 0.02 1.02 .± 0.03 1.16 .±. 0.03 1.14 .±. 0.04 1.16 ± 0.01 0.98 .± 0.01 1.07 ± 0.03

Tp59H 5889 .±. 148 48.8 .±. 0.2 0.82 .± 0.01 1.40 .± 0.04 3740 .± 40 31.7 .±. 1.6 45.2 .±. 1.0 8.1 .± 0.0 5.0 .±. 0.3 1.04 .±. 0.02 Tp60H 7745 .±. 122 627 ± 1.5 0.76 .±. 0.01 1.65 .±. 0.03 4694 .±. 64 43.6 .±. 1.1 55.7 .±. 1.1 4.8 .± 0.1 7.1 .±. 0.1 Tp61H 7297 .± 152 62.5 .±. 1.4 0.74 .±. 0.00 1.65 .±. 0.02 4107 .±. 64 39.0 .±. 0.5 53.1 .±. 1.4 4.8 ± 0.1 7.1 .± 0.3 Tp62H 6708 .± 144 56.5 .± 0.4 0.73 .±. 0.02 1.58 .±. 0.01 3758 .±. 86 35.3 .±. 2.3 48.9 .± 1.1 4.9 .± 0.2 7.1 .± 0.4 Tp63H 5907 .± 61 38.0 .± 0.1 0.54 .±. 0.02 1.20 .±. 0.01 3906 .±. 113 30.6 .± 0.5 41.3 .±. 0.5 3.6 .± 0.1 4.9 .± 0.2 Tp64H 5944 .± 42 32.5 .±. 0.2 0.56 .±. 0.01 1.23 .± 0.01 5092 .± 73 28.0 .±. 0.5 40.8 .± 0.8 4.1 .±. 0.0 4.6 .±. 0.2 Tp65H 5658 .± 60 40.9 ± 0.5 0.57 .±. 0.02 1.21 .±. 0.01 4151 .±. 34 30.5 .± 0.4 41.8 .± 0.7 4.2 .± 0.0 5.1 .± 0.1 Tp66H 7383 .±. 141 53.5 .±. 0.4 0.58 .± 0.01 1.33 .± 0.03 4719 .± 91 44.5 .± 1.7 59.2 .±. 1.0 5.5 ± 0.0 6.5 .± 0.2 Tp67H 7228 .± 328 57.7 .± 2.7 0.54 .± 0.02 1.25 .± 0.05 4660 .± 177 49.5 .± 2.7 61.2 .±. 3.3 6.7 .± 1.0 6.4 .± 0.5

=±= ~±~

~±=±= =±= =±= 1.19 .± 0.03 1.60 .± 0.38

~Tp68H 6707 .± 81 48.8 + 0.4 0.46 + 0.03 1.24 + 0.01 5676 + 33 34.7 ± 0.4 54.6 + 0.4 6.6 .±. 0.1 6.6 + 0.1 1.03 .± 0.02 @: for explanation o£ sample number refer to APPENDIX If.

• : n""2

'

.... u. "'

Appendix Illb: Selected elemental composition of termite mounds sampled at Daly River, site 3, following perchloric/nitric acid (4:1) extraction.

Sample Number@ Nt01D3• Nt02D3 Nt03D3• Nt04D3• Nt05D3 Nt06D3• Nt07D3

Species: Nasutitermes triodiae.

Element + sd (n=3) mgllOOg AI Ca Co Cu Fe

5139 .± 163 26.6 .± 0.2 0.52 .± 0.02 0.72 .± 0.03 3995 .± 125 4891 .± 86 20.1 .± 0.0 0.52 .± 0.02 0.73 .± 0.01 3633 .± 109 4514 .± 231 30.5 .± 0.4 0.54 .± 0.03 1.09 .± 0.06 3405 .± 31 4919 .±. 166 24.3 .± 0.2 0.54 4878 .:!: 22 23.0 4638 .± 211 17.7

.± 0.2

.± 0.3 4550 .± 292 22.0 .± 0.1

0.50 0.53 0.55

.± 0.00 0.77 .± 0.03 3566 .± 22

.± 0.01 1.03 .± 0.01 3629 .± 50

.± 0.00 1.12 .± 0.04 3461 .± 89

.:!: 0.01 0. 78 .:!: 0.02 3287 .:!: 135

K Mg 1039 .± 27 268.8 .± 3.7 937 .± 8 244.2 .± 2.9 1039 .:!: 19 247.7 .:!: 7.3 1023 .± 41 271.5 .± 3.8 1052 .± 50 246.0 .± 2.5 966 .± 52 237.6 .± 7.0 914 .± 39 246.6 .± 7.7

Nt08D3• 4400 .± 106 26.0 .± 0.3 0.50 .± 0.02 0.82 .± 0.02 3551 .± 90 870 .± 19 238.5 .± 4.1 Nt09D3 4770 .± 183 27.3 .± 1.2 0.51 .± 0.01 0.74 .± 0.03 3422 .± 189 859 .± 48 321.0 .± 12.6 Nt10D3 4421 .± 37 19.0 .± 0.2 0.57 .± 0.01 0.66 .± 0.00 3267 .± 103 783 .± 52 248.6 .± 2.1 Nt11D3 4458 .± 79 20.8 .± 0.3 0.51 .± 0.01 1.19 .± 0.06 3436 .± 114 941 .± 11 229.2 .± 3.7 Nt12D3 4567 .± 241 22.4 .± 0.1 0.49 .± 0.02 0.64 .± 0.01 3487 .± 34 939 .± 59 255.5 .± 6.3

Mn Na Zn 10.0 .± 0.0 19.9 .± 0.6 1.52 .± 0.06 8.5 .± 0.1 17.9 .± 0.2 1.54 .± 0.08 8.3 .± 0.3 21.0 .± 1.6 1.54 .± 0.12 12.8 .± 0.3 18.5 .± 1.1 1.68 .± 0.03 9.4 .± 0.2 18.7 .± 0.2 1.68 .± 0.06 8.7 .± 0.1 18.0 .± 1.3 1.74 .± 0.05 9.9 .± 0.1 15.7 .± 1.5 1.61 .± 0.05 9.2 .± 0.1 14.8 .± 0.9 1.63 .± 0.07 10.6 .± 0.3 20.2 .± 0.8 1.69 .± 0.05 8.0 .±. 0.2 13.7 .± 0.3 1.55 .± 0.02 9.3 .± 0.1 15.7 .± 0.8 1.65 .± 0.05 9.2 .± 0.0 16.2 .±. 1.6 1.51 .± 0.06

Nt13D3 4637 .± 95 18.7 .± 0.1 0.57 .± 0.01 1.32 .± 0.06 3639 .± 69 956 .± 49 267.3 .± 1.3 10.5 .± 0.2 16.3 .± 0.3 1.97 .± 1.59 .± 1.80 .± 1.69 .±

0.07 0.02 0.08 0.06

Nt14D3 4643 .± 48 30.2 .± 0.2 0.49 .± 0.01 0.83 .± 0.02 3386 .± 88 868 .± 55 298.8 .± 1.7 10.5 .± 0.1 20.0 .± 0.3 Nt15D3 4202 .± 166 27.3 .± 0.1 0.51 .± 0.02 1.20 .± 0.09 3387 .± :tS 893 .± 29 237.1 .± 3.1 8.8 .± 0.1 16.9 .± 1.0 Nt16D3# 4700 .± 48 17.6 .± 0.1 0.57 .± 0.01 1.17 .± 0.09 3352 .± 26 1075 .± 55 241.3 .± 1.6 7.8 .± 0.2 Nt17D3# 4422 .± 225 19.7 .± 0.3 0.54 .± 0.00 0.68 .± 0.01 3333 .± 56 912 .± 34 244.7 .± 6.2 7.6 .± 0.1 Nt18D3# 4588 .± 165 20.6 .± 0.4 0.59 .± 0.00 1.13 .± 0.02 3284 .± 96 947 .± 24 256.3 .± 4.6 8.8 .± 0.2

16.4 .± 0.5 13.7 .± 0.9 14.4 .:!: 0.9

1.58 .±. 0.05 1.73 .± 0.06

Nt19D3# 4102 .± 287 29.2 .± 0.6 0.56 .± 0.02 0.74 .± 0.02 3008 .± 137 817 .± 39 261.6 .± 8.5 10.9 .± 0.5 13.2 .± 0.9 1.62 .±. 0.05 Nt20D3## 4590 + 63 73.6 + 0.5 0.59 + 0.01 1.18 + 0.01 3248 + 103 991 + 56 297.8 + 2.5 13.5 + 0.2 15.3 + 0.2 2.10 + 0.01

@; for explanation of sample number refer to APPENDIX Ig.

... ..,. ....

Appendix lllh: Selected elemental composition of termite mounds sampled at Daly River, site 4, following pen:hloric/nitric acid (4:1) extraction.

Sample Number@ Nt21D4* Nt22D4 Nt23D4 Nt24D4

• Nt25D4 " Nt26D4

Nt27D4 Nt28D4 Nt29D4 Nt30D4* Nt31D4* Nt32D4* Nt33D4 Nt34D4 Nt35D4

" Nt36D4* Nt37D4 Nt38D4 Nt39D4 Nt40D4*

" Nt41D4* Nt42D4* Nt43D4 Nt44D4 Nt45D4

AI Ca 4817 .± 136 33.0 .± 0.8 0.37 4870 .± 139 3592 .± 19

40.0 .± 0.2 30.3 .± 0.5

4610 .± 222 28.3 + 0. 4

0.37 0.31 1131 0.35 0.34 0.25

4496 .± 6 4531 .± 119 4008 .± 77 4330 .± 78

49.5 .± 0.2 34.7 32.6

.± 0.7

.± 0.3 25.2 .± 0.6 0.26

Co + 0.02 .± 0.01

Species: Nasutitermes triodiae.

Element + sd (n=3) mg/1()0g

0.81 0.84

Cu Fe K .± 0.02 1374 .± 25 819 .± 17 .± 0.01 1408 .± 4 ~66 836

.± 0.00 0.75 .± 0.01 .± 0.03 .± 0.00 .± 0.04 .± 0.01 .± 0.01

1350 .± 17 593 .± 17 .± 0.03 .± 0.01 .± 0.01 .± 0.00 .± 0.01

0.92 0.87 0.87 0.75 0.81

2165 .± 50 620 .± 24 1714' .± 1700 .±

22 663 .± 35 0 707 .± 28

.± 5 1271 .± 8 614 1621 .±

3580 .± 72 52.8 .± 0.2 0.26 .± 0.01 0.75 .± 0.01 1264 .± 20 582 37 44.8 .± 0.7 0.27 .± 0.01 0.76 .± 0.00 1279 .± 10 585

.± 0.2 0.27 .± 0.01 0.72 .± 0.01 1241 .± 5 554 3708 .± 3478 .± 22

68 43

35.3

11 608 .± 9 .± 14 .± 8 .± 3 .t 17 2939 .±

4008 .± 4111 .± 3479 .±

29.6 .± 0.4 0.22 .± 0.01 0.60 .± 0.01 60.7 .± 0.7 0.30 .± 0.01 0.87 .± 0.02

25 33.7 .± 0.3 0.30 .± 0.00 0.87 .± 0.01 77 37.4 .± 0.9 .± 0.02

3676 .± 110 24.1 .± 0.2 0.28 0.30 .± 0.01

0.78 0.78

.± 0.03

.± 0.02

961 1226

.± 7 ~ 4

1240 .± 12 1063 .± 12 1182 .t13

505 656 .± 28 738 .± 21 672 699

3188 .± 53 3432 .± 125 3381 .± 63

18.7 .± 0.3 16.7 .± 0.3

0.23 0.23 0.24

.± 0.00 0.69 .± 0.00 947 .± 6 515

.t 15

.± 38

.± 21

.± 41

.± 30

.± 11

.± 2

.± 32

.± 22

22.2 2867 .± 37 33.5

+ 0.4 .± 0.4

.t 0.02

.± 0.01 0.22 .± 0.01

2850 .± 10 28.7 .t 0.1 0.22 .± 0.02 3286 .± 16 25.8 .± 0.4 0.24 .± 0.01 3816 .± 135 49.9 .± 1.6 0.28 .± 0.02 3728 .± 20 41.4 .± 1.1 0.30 .± 0.02 3986 .± 23 36.7 .± 0.8 0.30 .± 0.02

0.74 .± O.o! 0.75 .± 0.01 0.68 0.59 0.71 0.76 0.69

.± 0.01

.± 0.00

.± 0.01

.± 0.05

.± 0.01

1044 .± 23 578 1238 .± 2 567

488 924 .± 7 .t 4 567

987 .± 7 590 1393 .±

799

48 675 580 -t 26 1184 .± 36

0.73 .± 0.01 1355 .± 11 651 .± 19

'

Mg Mn Na .t 1.7 3.2 .± 0.1 28.3 93.7

103.8 79.0

.± 1.5 3.7 .t 0.1 30.7 .± 0.5 .± 1.8 .t 1.7

0.41 0.41 0.39 .± 1.0 3.8

26 102.4 .± 5.8 .± 0.0 .± 0.0

22.4 23.3 .± 22 0.47 37.5 .t 0.8 0.39

Zn .± 0.01 .± 0.01 .± 0.01 .t 0.03 .± 0.04 134.6 .± 0.5

111.8 .± 4.1 4.4 .± 0.1 3.5 .± 0.1 29.8

21.8 .t 1.6 .± 1.1

0.47 .± 0.01 .± 1.7 2.3 .± 0.0 .± 0.9 1.9 .± 0.0 21.5 .± 1.8

0.46 0.53

.± 1.8 3.4 .± 0.1 22.9 .± 1.8 0.45

.± 0.4 3.8 .t 0.0 21.5

.± 0.1 3.0 .± 0.0 17.4

.± 1.1 2.8 4.7 .± 0.9

.± 0.7

.± 0.2 0.46 0.44

.t 1.5 0.38

.± 0.6 0.43

.± 0.04

.± 0.08

.± 0.03

.± 0.01

.± 0.03

.± 0.04

.± 0.01

85.5 82.2 %.2 103.0 85.8 87.1 118.7 89.7 .± 0.4 3.5

3.2

.± 0.0

.± 0.0

.± 0.0

.± 0.0

20.8 26.5 30.0 29.8 28.6

.t 0.1

.± 1.2

.± 1.0

.t 1.2

0.44 .± 0.03 130.1 .t 27

86.4 3.5 .t 0.1 67.2 67.6 72.4 85.0 93.5

.± 1.3

.± 1.1 1.5 .± 0.0 15.8

0.39 0.39 0.38

.± 2.4 1.4

.± 2.4 1.6 .t 0.0 .± 0.0

.t 0.8 2.5 .t 0.0

.± 0.4 2.4 .± 0.0 81.4 .± 0.6 2.2 .± 0.0 100.6 91.7

.± 3.4

.± 1.2 4.4 .± 0.3 3.3 .± 0.1

18.1 .± 1.9 0.40 20.3 .± 2.0 0.38 17.8 23.2

.± 0.5 0.42

.± 0.1 21.1 .± 0.2 26.4 22.4

96.5 .t 0.8 3.5 .± 0.0 27.6

.± 0.4

.t 0.9

.t 0.5

0.37 0.42 0.51 0.43 0.45

.±~

.±M1

.±=

.±~

.±~

.±=

.±=

.±=

.±~

.±~

.±M1 continued •. ,

~

APPENDIX Illh: continued

Sample Number@ Nt46D4 Nt47D4 Nt48D4 Nt49D4* Nt50D4* Nt51D4* Nt52D4 Nt53D4 Nt54D4 Nt55D4 Nt56D4 Nt57D4 Nt58D4* Nt59D4* Nt60D4* Nt61D4* Nt62D4 Nt63D4 Nt64D4 Nt65D4 Nt66D4 Nt67D4* Nt68D4* Nt69D4* Nt70D4* Nt71D4

~ Nt72D4

AI Ca 3305 .±. 126 3366 .± 30 3804 ± 48

20.3 .±. 0.5 31.0 33.4

3339 .±. 64 33.0 3511 ± 24 49.1

26.3 23.5

+ 0.5 .± 1.2 + 0.5 .±. 0.7 .±. 0.2 .± 0.4

0.21 0.20 0.23 0.21 0.23 0.19 0.39

Element + sd (n=3) mg/100g Co Cu Fe K + 0.01 .± 0.01

0.65 .±. 0.02 1176 .± 23 563 .± 39 0.66 .± 0.00 1052 .±. 2 608 .±. 24

.±. 0.01 0.73 .±. 0.00 1262 .± 6 623 .±. 15 .± 0.00 1075 .± 9

± 4 1109 603 .± 22 619 .± 6 581 .± 33 689 .± 47

3075 .± 14 5300 .± 43 5925 ± 113 4331 .± 25 4645 .± 33 4334 .± 89 4547 .± 100 3537 .±. 65

21.8 .± 0.1 0.43

+ 0.00 .± 0.01 .± 0.00 .± 0.02 .± 0.01 .± 0.00 .± 0.01

0.66 0.71 0.61 0.90 1.02 0.78 0.90

.± 0.00

.± 0.01

.± 0.01

.± 0.01

.± 0.01

966 .± 9 1538 .± 6 2128 .± 1518 .±

34 722 .± 23 3 596 .± 47

.± 22 ± 5

22

27.6 25.8 34.0 39.2

.± 0.3

.± 0.3

.± 0.6

.± 0.5 26.4 .± 0.1

0.36 0.32 0.31 0.32 0.26

.± 0.00 1407 .± 10 621 .± 0.01 0.88 .± 0.01 1473 .± 16 618 .± 0.00 .±. 0.01

0.91 .± 0.04 1544 .±. 17 0.73 .±. 0.02 1018 ± 5

633 .±. 546 .± 19

3317 .±. 66 25.8 .±. 0.1 0.25 .±. 0.00 0.69 .±. 0.01 1095 .±. 7 552 .± 9 3224 .± 58 27.2 .±. 0.3 0.25 .± 0.00 0.72 .±. 0.00 1007 .± 4 508 .±. 13 4257 .±. 16 11.9 .±. 0.2 0.41 .± 0.01 0.76 .±. 0.00 1303 .± 8 623 .±. 13 3787 .±. 40 23.7 3844 .± 193 22.9

.±. 0.4

.± 0.6 0.30 + 0.01 0.76 0.33

1540 .± 15 662 .±. 7 1649 ± 13' 667 ± 9

5073 .±. 124 4514 .± 151 3279 .± 34 3547 .± 31 3351 .±. 6 2212 .±. 60 4042 .± 67 4323 .±. 21 3505 .± 31

16.0 .±. 0.2 0.41 + 0.00 .± 0.02

0.80 0.97

+ 0.02 .± 0.02 .±. 0.00 .± 0.02

2252 ± 30 ±25

.±. 0.01 1060 .±. 5

646 .±. 15 641 .± 29 617 ± 9 626 .±. 19 630 .± 5

18.6 16.8

+ 0.6 .±. 0.2

20.8 .± 0.0 18.6 .±. 0.2 13.4 .±. 0.1 16.2 28.4

+ 0.3 .±. 0.2

36.4 .±. 0.1

0.37 .± 0.01 0.85 1839 0.26 0.30 0.30 0.21 0.36 0.39

.± 0.01

.±. 0.01 + 0.02 .±. 0.01 + 0.01 .±. 0.04

0.32 + 0.01

.±. 0.01 1344 .±. 13 1155 .±. 14

0.68 0.70 0.65 0.51

+ 0.01 .±. 0.00 899 .±. 2

0.84 .± 0.01 2125 493 .±. 33

.± 11 749 .±. 11 0.88 .±. 0.01 2093 .±. 9 0.79 + 0.02 1673 ± 5

769 ± 14 662 .±. 13

Mg Mn 73.6 ± 0.1 79.1 .±. 0.2

.±. 2.3 1.7 2.2 .± 0.0 2.4 .± 0.0

17.8 20.1 22.4 20.8 24.7 20.0 32.1

104.1 81.5 110.1 77.3 111.7 126.9 91.6 112.1

.±. 1.4

.±1.1 2.8 ± 0.0 ± 0.4 3.9 .± 0.4 23

+ 0.0 .±. 0.0

± 1.3 2.8 .±. 0.1

.± 0.6

.±. 0.4

.± 1.2 2.8 ± 0.0 29.5 3.4 + 0.0 24.0 2.3 + 0.0 25.2 2.8 ± 0.0 23.1 109.5 ± 1.0

150.9 ± 2.3 3.4 .± 0.0 30.0 27 .±. 0.0 21.7

Na

.±. 2.3 0.45

.± 0.5 0.47

.±. 0.4

.±. 1.2 0.47 0.45

± 0.6 0.48

Zn ± 0.05 .±. 0.01 .± 0.00 .±. 0.01 .±. 0.01

.± 0.1 0.40 .±. 0.01

.±. 0.6

.±. 0.3 0.59 .±. 0.15 0.49 .±. 0.02

.± 0.1 0.40 .± 0.01

.±. 0.9 0.52 .±. 0.1 0.49

.± 0.02 ± 0.01

.±. 1.2

.±. 1.3 0.50 .±. 0.04

%.1 88.9

+ 0.9 .± 0.5 2.9 .±. 0.0 19.4 .±. 0.4

0.44 0.46 0.52

.±. 0.03

.± 0.04

.± 0.04

.± 0.02 85.0 .± 1.1 2.7 ± 0.0 17.2 .± 0.7 94.3 .±. 1.9 2.2

3.1 .±. 0.0 .± 0.1

26.8 .±. 0.6 0.35 102.3 ± 0.1 109.0 .±. 1.8 2.9 .± 0.0

27.7 28.5

125.3 .± 2.5 1.6 ± 0.0 30.1 113.6 .±. 1.4 1.7 .±. 0.0 27.9 87.3 .±. 0.4 1.5 ± 0.0 23.4 97.3 .±. 1.2 1.8 .± 0.0 26.1 98.2 ± 1.6 1.8 .±. 0.0 27.6 69.4 .±. 0.7 1.3 .± 0.0 20.0 110.0 .± 2.7 2.5 .± 0.0 30.5 133.8 .± 0.3' 3.1 .±. 0.0 33.0 121.3 ± 0.9 3.6 .±. 0.0 30.7

.±. 0.7 0.40 .±. 0.01

.±. 0.7 .±. 1.3

0.38 0.51

± 0.00 .±. 0.03

.± 0.5 0.45 .±. 0.02

.±. 0.5 0.36 .± 0.03 ± 0.4 0.40 .±. 0.01 .±. 0.6 0.38

0.28 .±. 0.00 ± 0.01 .±. 1.1

.±. 0.7 0.41 ± 0.01 ± 0.2 0.46 .±. 0.00 .±. 0.6 0.40 .± 0.01

continued ... "' "' "'

APPENDIX lllh: continued ...

Sample Number@ Nt73D4 Nt74D4 Nt75D4 Nt76D4 Nt77D4 Nt78D4 Nt79D4 Nt80D4 Nt81D4

" Nt82D4 Nt83D4 Nt84D4 Nt85D4* Nt86D4 Nt87D4 Nt88D4 Nt89D4 Nt90D4 Nt91D4

Element + sd _(n=3) mgl100g AI Ca Co Cu Fe K Mg Mn Na Zn

3751 ± 44 38.6 ± 0.8 0.29 ± 0.01 0.90 ± 0.03 1479 ± 5 627 ± 28 97.8 ± 0.8 3.4 ± 0.0 22.8 ± 1.9 0.51 .± 0.04 3654 ± 35 33.3 ± 0.5 0.30 ± 0.01 0.85 ± 0.01 1423 ± 9 648 ± 9 92.3 ± 0.7 3.5 ± 0.0 24.0 ± 0.5 0.43 .± 0.00 3419 ± 26 24.5 ± 0.3 0.31 .± 0.01 0.76 .± 0.01 1204 ± 26 619 ± 12 83.1 .± 1.1 28 ± 0.0 23.4 .± 0.0 0.47 ± 0.00 4074 ± 48 78.9 ± 1.7 0.35 ± 0.01 0.93 ± 0.02 1538 ± 33 636 ± 37 130.5 ± 2.9 6.6 ± 0.1 21.1 ± 0.6 0.61 ± 0.08 3821 ± 59 104.0 ± 0.6 0.32 ± 0.02 0.90 ± 0.00 1511 ± 11 570 ± 44 146.4 .± 0.6 6.9 ± 0.0 24.8 ± 0.5 0.54 ± 0.03 3942 ± 82 68.8 ± 1.6 0.36 ± 0.00 0.91 .± 0.02 1552 .± 10 657 ± 24 126.5 ± 1.3 5.9 ± 0.0 25.3 .± 2.2 0.48 ± 0.03 4479 ± 100 93.4 ± 2.4 0.38 .± 0.01 1.00 ± 0.01 1684' ± 9 780 .± 18 126.7 ± 2.2 5.0 ± 0.1 25.6 ± 0.3 0.62 .± 0.02 4554 ± 95 79.5 ± 1.0 0.39 .± 0.01 1.01 ± 0.02 1765 ± 8 762 ± 39 125.9 ± 1.3 5.5 ± 0.1 26.5 ± 0.7 0.63 ± 0.02 4426 .± 35 116.4 .± 1.4 0.37 .± 0.01 0.90 .± 0.01 1633 .± 23 653 .± 24 160.0 .± 21 3.8 .± 0.0 35.1 .± 0.6 0.56 .± 0.03 4722 .± 30 23.5 ± 0.6 0.32 ± 0.01 0.98 .± 0.00 1740 .± 23 651 ± 14 99.1 ± 0.1 2.7 ± 0.0 220 .± 0.0 0.45 ± 0.01 3662 ± 39 30.4 .± 0.4 0.29 .± 0.01 0.76 ± 0.01 1288 .± 3 596 .± 27 91.7 .± 0.7 3.1 .± 0.1 22.1 ± 0.4 0.38 .± 0.01 3165 ± 68 51.7 ± 0.5 0.33 .± 0.01 0.70 ± 0.00 998 ± 13 562 .± 17 90.9 .± 0.7 4.3 ± 0.1 22.8 ± 0.4 0.39 .± 0.01 3322 .± 90 32.1 ± 0.2 0.33 .± 0.03 0.71 ± 0.01 1047 ± 5 572 ± 30 83.5 ± 1.2 3.7 ± 0.0 23.3 ± 1.2 0.37 ± 0.01 3820 .± 72 29.6 .± 0.2 0.34 ± 0.01 0.78 .± 0.01 1326 ± 3 640 ± 23 88.1 ± 0.6 3.3 .± 0.1 22.5 .± 0.5 0.42 .± 0.02 3479 ± 25 29.2 .± 0.3 0.32 .± 0.01 0.70 .± 0.01 1077 .± 2 593 .± 14 85.2 .± 0.6 3.4 .± 0.1 22.3 .± 0.3 0.35 .± o.oo 3606 ± 58 40.6 ± 0.7 0.33 .± 0.01 0.77 ± 0.01 1152 ± 23 610 .± 34 95.7 .± 0.5 3.7 .± 0.1 26.1 .± 0.7 0.41 .± 0.01 3248 .± 29 32.8 .± 0.2 0.30 .± 0.01 0.69 .± 0.01 1052 .± 3 555 .± 26 86.4 .± 0.8 3.4 .± 0.1 23.9 .± 0.8 0.41 .± 0.02 3891 .± 66 40.8 .± 0.5 0.33 .± 0.01 0.82 .± 0.02 1404 .± 9 658 .± 18 91.2 .± 1.5 3.1 .± 0.0 24.7 .± 0.7 0.43 .± 0.03 3528 .± 1~- 29.1 ± __ Q.3 0.31 + 0.02 0.83 + 0.01 1364 + 3 645 + 7 92.4 + 0.8 3.3 .± 0.0 __ 26.4 + 0.3 0.41 .± 0.00

@:for explanation of &ample number refer to APPENDIX lh.

": n=2.

~ <:>

Appendix Illi: Selected elemental composition or termite mounds sampled at Howard Springs (site 6) and Berrimah (site 7) rollowing

Sample Number@

Nt92H Nt93H* Nt94H Nt95H Nt%H Nt97H Nt98H Nt99H NtlOOH Nt101H Nt102H Nt103H Nt104H Nt105H Nt106H

Nt107B Nt108B Nt109B NtllOB Nt111B

perchloric/nitric acid (4:1) extraction. Species: Nasutiternres lriodiae.

Element .± sd (n=3) mg!lOOg AI Ca Co Cu Fe K Mg Mn

4834 .± 47 60.6 .± 0.6 0.56 .± 0.00 1.35 .± 0.02 3283 .± 32 34.3 .± 1.3 40.6 .± 0.6 3.2 .± 0.1 4725 .± 34 44.6 .± 0.5 0.54 .± 0.03 1.32 .± 0.00 2796 .± 31 30.1 .± 1.6 37.7 .± 0.4 2.8 .± 0.0 4636 .± 40 57.3 .± 0.2 0.54 .± 0.02 1.29 .± 0.01 2475 .± 18 32.5 .± 0.7 43.0 .± 0.5 2.9 .± 0.0 5141 .± 9 82.4 .± 0.3 0.65 .± 0.01 1.54 .± 0.01 3289 .± 41 36.3 .± 0.3 48.6 .± 0.3 5.2 .± 0.1 4960 .± 116 66.4 .± 1.7 0.62 .± 0.02 1.50 .± 0.03 3019 .± 42 39.5 .± 0.6 43.7 .± 0.7 5.0 .± 0.1 5622 .± 56 56.9 .± 0.2 0.70 .± 0.02 1.61 .± 0.01 3300 .± 50 38.7 .± 0.5 46.0 .± 0.5 4.3 .± 0.1 5949 .± 61 80.0 .± 1.0 0.69 .± 0.01 1.54 .± 0.00 3739 .± 147 35.6 .± 0.2 48.5 .± 0.1 3.7 .± 0.0 5703 .± 37 88.1 .± 1.1 0.68 .± 0.03 1.51 .± 0.02 3562 .± 69 35.8 .± 0.5 48.4 .± 0.7 4.0 .± 0.1 5638 .± 28 80.7 .± 0.4 0.67 .± 0.02 1.50 .± 0.00 2927 .± 71 36.7 .± 1.5 48.8 .± 0.1 3.7 .± 0.0 6364 .± 49 113.9 .± 1.1 1.03 .± 0.01 1.69 .± 0.01 3868 .± 132 41.2 .± 0.3 57.2 .± 1.0 5.8 .± 0.0 6509 .± 102 100.3 .± 3.1 1.00 ± 0.02 1.66 ± 0.02 4056 ± 49 41.9 ± 0.5 54.8 .± 1.2 5.9 ± 0.0 6485 ± 154 120.9 .± 3.1 1.03 .± 0.02 1.67 .± 0.02 4205 .± 21 47.0 .± 4.9 64.7 ± 0.9 7.0 ± 0.1 6584 ± 380 89.6 .± 4.3 1.11 .± 0.03 1.93 .± 0.26 4236 .± 102 42.8 .± 5.7 55.8 .± 2.8 6.5 .± 0.1 6717 .± 66 111.4 .± 2.0 1.08 .± 0.01 1.71 .± 0.00 4687 .± 52 44.0 .± 2.9 59.2 .± 1.1 7.7 .± 0.1 6200 .± 85 101.8 ± 0.9 0.95 ± 0.01 1.59 .± 0.01 5288 .± 74 41.1 .± 2.0 57.5 .± 0.8 7.8 .± 0.1

Na Zn 4.9 ± 0.1 0.90 .± 0.01 4.8 .± 0.3 0.67 .± 0.18

0.02 0.02 0.04 0.02 0.05

6.7 .± 0.1 6.5 .± 0.1 6.4 .± 0.0 5.6 .± 0.1 5.4 .± 0.3 5.1 .± 0.2 5.4 .± 0.1 5.9 .± 0.1

0.79 .± 1.14 .± 0.96 ± 1.03 .± 1.17 ± 1.11 0.98 1.21

.± 0.05

.± 0.01

.± 0.01 5.9 .± 0.3 7.9 .± 0.0

1.17 .± 0.04 1.09 .± 0.02

6.0 .± 0.2 1.20 .± 0.05 6.0 .± 0.1 1.18 .± 0.02 5.9 .± 0.1 1.02 .± 0.01

2999 ± 94 97.5 ± 1.9 0.37 .± 0.01 0.83 ± 0.02 882 .± 12 3741 ± 106 78.2 .± 1.0 0.36 .± 0.01 0.93 ± 0.02 1121 .± 14 2972 ± 34 122.5 .± 0.9 0.31 .± 0.00 0.87 .± 0.02 842 .± 19 3412 .± 94 53.3 .± 0.7 0.32 .± 0.01 0.88 .± 0.02 823 .± 11 3529 .± 68 60.3 .± 1.3 0.33 .± 0.01 0.93 .± 0.01 782 .± 5

317 .± 19 65.9 .± 2.0 3.5 .± 0.1 35.3 ± 2.2 369 .± 21 58.5 .± 1.1 3.0 .± 0.1 32.5 .± 1.7 335 .± 4 63.0 .± 0.6 4.0 .± 0.0 38.5 .± 0.7 335 ± 24 52.5 .± 1.8 2.6 .± 0.1 28.7 .± 1.8 347 .± 4 53.9 .± 0.9, 2.9 .± 0.1 32.1 .± 1.5

0.57 .± 0.04 0.68 .± 0.04 0.67 .± 0.02 0.69 + 0.04 0.61 .± 0.01

@:for explanation of sample number refer to APPENDIX Ii. ... "' ~

... i:l

Appendix Illj: Selected elemental composition or termite mounds sampled at Daly River, site 1, rollowing pen:hloric/nitric acid (4:1) extraction. Species: (Tumulitermes hastilis)

Sample Element + sd {n-3~ mgL100g

Number@ AI Ca 0> Cu Fe K Mg Mn Na Zn

Th01D1 2241 .± 25 92.3 .± 3.3 0.32 .± 0.02 0.55 .± 0.01 1297 .± 12 419 ± 12 77.5 ± 1.2 6.5 + 0.0 19.1 ± 1.7 0.42 ± 0.01

Th02D1 2570 + 48 79.8 .± 1.3 0.37 + 0.02 0.69 .± 0.01 1406 .± 115 382 ± 6 70.5 ± 0.8 5.3 .± 0.1 17.9 ± 0.5 0.52 ± 0.01

Th03D1 2280 + 75 34.2 .± 0.7 0.28 .± 0.01 0.50 .± 0.01 1259 .± 10 419 ± 12 57.7 ± 1.4 3.7 .± 0.0 19.8 ± 1.2 0.37 ± 0.01

Th04D1 1969 .± 29 120.2 .± 0.5 0.26 .± 0.01 0.52 .± 0.01 1118 .± 4 384 ± 16 79.5 ± 0.5 5.8 .± 0.1 17.4 .± 0.1 0.35 ± 0.00

Th05DI 1986 .± 44 212.9 .± 3.5 0.28 + 0.01 0.58 . .± 0.01 1166 .± 18 410 ± 22 96.5 ± 0.7 8.3 .± 0.1 17.8 ± 1.2 0.42 ± 0.00

@; for explanation of sample number refer to APPENDIX lj.

APPENDIX lllk: Selected elemental composition or soils (0·10cm depth) sampled at Elliott (siteS), Daly River (1-4), Howard Springs (site 6)

and Berrimab (site 7)1 rollowing perchloric/nitrlc acid (4:1) extraction.

Sample Element .± sd (n=3) mgflOOg Number AI Ca Co Cu Fe K Mg Mn Na Zn

OlE 1614 .± 62 45.6 .± 1.3 0.26 .± 0.03 0.62 .± 0.03 1063 .± 36 88.3 .± 5.2 44.4 .± 1.0 5.90 .± 0.27 4.04 .± 0.27 0.51 .± 0.05 02E 2080 .± 21 48.9 .t 0.6 0.27 .± 0.01 0.74 .± 0.01 1244 .± 13 105.1 .± 2.7 52.2 .t 0.5 6.83 .t 0.18 4.02 .± 0.18 0.66 .± 0.04 03E 1981 .t 5 69.1 .± 0.7 0.32 .t 0.04 0.72 .± 0.03 1208 .± 7 113.1 .± 0.9 54.9 .± 0.5 12.49 .± 0.05 3.98 .± 0.05 0.79 .t 0.06 04E 2559 .± 81 40.8 .t 0.9 0.24 .± 0.02 0.76 .± 0.01 1409 .± 30 108.0 .± 3.3 62.4 .± 1.5 3.88 .± 0.12 4.41 .± 0.25 0.80 .± 0.04

0501 1713 .± 81 7.0 .± 0.3 0.20 .± 0.01 0.39 .± o.oo 957 .± 8 367.4 .± 23.4 31.6 .± 1.9 3.19 .± 0.06 17.32 .± 2.46 0.39 .± 0.01 0601 1838 .± 100 5.4 .± 0.0 0.19 .± 0.01 0.41 .± 0.01 1007 .± 14 421.9 .± 19.1 32.0 .± 1.5 2.39 .± 0.05 19.55 .± 1.91 0.37 .± 0.02 0701 1617 .±. 92 10.7 .±. 03 0.19 .±. 0.01 0.37 .±. 0.00 1070 .±. 8 330.1 .±. 3.5 30.8 .±. 1.1 3.53 .±. 0.07 14.45 .±. 1.69 0.40 ±. 0.01 0801 1663 .± 44 226.5 .± 5.5 0.25 .± 0.00 0.61 .± 0.00 1078 ± 16 388.4 .± 8.2 71.3 .± 0.4 9.29 .± 0.10 15.91 .± 0.72 0.47 .± 0.03 0902 2642 .± 68 11.3 .± 0.0 0.45 .± 0.02 0.68 ± 0.01 1184 .± 19 379.9 .± 10.9 92.9 .± 1.2 3.86 .± 0.10 8.81 .± 0.41 0.77 .± 0.02 1002 3058 .± 72 5.2 .± 0.1 0.47 .± 0.03 0.69 .± 0.01 1200 .± 17 404.4 .± 17.4 93.1 .± 1.7 2.97 .± 0.10 9.60 .± 0.61 0.74 .± 0.01 1102 2842 .± 97 4.1 .± 0.1 0.41 .± 0.01 0.60 .± 0.00 1063 .± 8 385.1 .± 30.0 80.3 .± 2.8 2.41 .± 0.10 9.12 .± 0.29 0.64 .± 0.02 1503 2405 .± 41 12.7 .± 0.3 0.40 .± 0.02 0.70 .± 0.03 2924 .± 76 545.5 .± 34.0 93.0 .± 2.6 10.14 .± 0.17 13.15 .± 0.23 1.18 .± 0.04 1603 2109 .± 108 14.0 .± 0.5 0.33 .± 0.04 0.73 .± 0.01 2725 .± 89 492.6 l 27.3 78.1 .± 2.8 8.88 .± 0.33 12.83 .± 0.56 4.83 .± 0.02 1703 3904 .±. 83 12.9 .± 0.3 0.51 .± 0.01 0.53 .± 0.00 2842 .± 10 942.9 .± 33.6 200.8 .± 1.7 6.62 .± 0.16 14.17 .± 0.26 1.63 .± 0.07 1803 3464 .± 211 5.2 .± 0.0 0.47 .± 0.01 0.49 .± 0.00 2671 .± 64 729.1 .± 49.9 165.8 .± 5.6 5.20 .± 0.08 10.39 .± 1.21 1.82 .± 0.39 1903 4057 .± 85 4.7 .± 0.1 0.53 .± 0.00 0.76 .± 0.01 3134 .± 123 859.9 .± 18.1 193.5 .± 1.5 5.53 .± 0.24 12.13 .± 0.14 1.50 .± 0.07 2003 2515 .± 32 10.1 .± 0.1 0.42 .± 0.02 0.67 .± 0.02 2976 .± 32 577.3 .± 13.6 93.7 .± 1.3 8.56 .± 0.13 13.88 .± 0.17 7.73 .± 0.52 2103 3340 .± 56 5.9 .± 0.1 0.47 .± 0.02 0.70 .± 0.01 2853 .± 45 643.8 .± 50.4 175.0 .± 2.8 5.64 .± 0.12 10.06 .± 0.63 1.46 .± 0.03 2203 3697 .± 151 11.0 .± 0.3 0.45 .± 0.02 0.50 .± 0.01 2347 .± 47 762.2 .± 50.6 157.7 .± 5.2 5.73 .± 0.08 14.12 .± 0.54 1.09 .± 0.07

continued ...

~

APPENDIX lllb: continued •••

Sample Element + sd {n=3) mg/100g Number AI Ca Co Cu Fe K Mg Mn Na Zn

2304 2759 .±. 17 29.7 .±. 0.8 0.30 .±. 0.01 0.61 .±. 0.01 1076 .±. 28 576.8 .±. 19.4 75.2 .±. 0.7 4.11 .± 0.02 20.78 .±. 0.75 0.41 .± 0.00 2404 3208 .±. 180 6.5 .±. 0.3 0.28 .± 0.01 0.71 .±. 0.02 857 .±. 11 652.6 .±. 39.8 67.2 .±. 3.8 2.36 .± 0.06 23.61 .±. 2.63 0.41 .± 0.01 2504 2119 .±. 41 6.5 .±. 0.3 0.23 .±. 0.01 0.43 .±. 0.01 570 .±.' 10 437.8 .±. 13.4 48.4 .± 0.8 1.66 .±. 0.03 16.27 .± 0.83 0.20 .±. 0.01 2604 2107 .±. 25 5.6 .±. 0.1 0.22 .±. 0.01 0.45 .±. 0.00 701 .±. 6 448.4 .±. 5.4 47.6 .±. 0.2 2.24 .±. 0.02 17.01 .±. 0.15 0.22 .±. 0.00 27H 4406 .±. 19 37.4 .± 0.1 0.44 .±. 0.02 1.22 .± 0.01 5114 .± 91 50.3 .±. 1.9 36.7 .±. 0.1 5.49 .± 0.07 4.10 .±. 0.11 1.05 .±. 0.00 28H 5508 .±. 47 101.8 .±. 0.9 0.97 .±. 0.01 1.51 .±. 0.01 5783 .±. 66 28.1 .± 2.2 45.9 .±. 0.3 12.88 .±. 0.21 4.32 .±. 0.16 2.59 .±. 0.22 29H 3789 .±. 29 53.6 .±. 0.5 0.77 .±. 0.01 1.31 .±. 0.02 7371 .±. 187 25.5 .± 0.2 31.2 .±. 0.1 11.50 .±. 0.06 3.77 .±. 0.07 2.69 .±. 0.03 30H 4472 .±. 48 34.4 .±. 0.1 0.46 .±. 0.01 1.09 .±. 0.01 5815 .±. 49 24.0 .±. 0.5 36.3 .±. 0.4 6.32 .±. 0.06 3.75 .± 0.04 1.07 .± 0.03 31B 2711 .± 58 24.9 .± 0.3 0.26 .± 0.01 0.92 .±. 0.01 3223 .± 7 292.9 .±. 17.2 34.4 .±. 1.0 2.11 .±. 0.11 18.62 .±. 0.71 0.56 .±. 0.01 32B 1553 .±. 14 16.0 .±. 0.2 0.27 .±. 0.00 0.46 .±. 0.00 599 .± 8 197.3 .±. 2.6 23.2 .±. 0.2 2.<J7 .±. 0.02 11.72 .±. 0.13 0.33 .±. 0.01 33B 1385 + 24 10.6 + 0.4 0.24 .± 0.02 0.40 .± 0.01 505 .±. 7 178.0 .± 8.9 20.5 .±. 0.2 1.95 .±. 0.08 10.35 .± 0.35 0.24 + 0.02

For selected termite species studied at each site, refer to APPENDIX Ik.

... ~

APPENDIX lVa: Selected elemental composition of termite mouods sampled at Elliott (site 6) and Daly River (2 & 4) following 0.1 N pepsin- IICI (pll 1.35) extraction

on lmm fraction size samples.

Species: Amitermes vitioms.


Number@ AI C• Co Cu Fo K Mg Mn N• Zn Fe{II)

Av01E 20.8 .± 0.2 128~ .±1.5 0.09 .± 0.01 0.20 .± 0.00 17.8 ± 0.3 20.6 .± 0.6 26.5 + 0.4 5.24 .± 0.14 1.16 .± 0.23 0.16 .± 0.16 13.3 .± 0.3

Av15E 20.3 .± 0.1 101.4 .± 0.9 0.06 .± 0.01 0.18 .± 0.00 16.0 .± 0.1 21~ .± 0.4 34.1 .± 0.3 2.60 .± 0.04 2.28 .± 0.15 0.06 .± 0.01 10.6 .± 0.2

Aill!l 21.1 .± 0.4 126.6 .± 3.1 0.06 .± 0.01 0.16 .± 0.00 20.9 .± Q3 16.0 .± 0.4 25.5 .± 0.4 3.84 .± 0.06 0.77 .± 0.05 0.09 .± 0.00 15.4 .± Q7

Av31D2 67.7 .± 0.8 32.6 .± 0.5 0.13 .± 0.02 0.17 .± 0.00 154.9 .± 3.7 14.4 .± 0.6 13.1 .± 0.4 5.58 .± 0.17 3.99 .± 0.19 0.20 .± 0.02 78.7 .± 0.9

Av34D2 46.7 .± 0.8 15.3 .± 0.4 0.05 .± 0.01 0.11 .± 0.00 66.1 ± 1.5 7.1 .± 0.4 5.2 .± 0.0 1.91 .± 0.02 1.97 .± 0.16 0.13 .± 0.02 36.3 .± 21

Av39D2 50.5 .± 1.4 24.6 .± 1.0 0.09 .± 0.00 0.13 .± 0.01 117.1 .± 24 7.5 .± 0.5 8.8 .± 0.2 4.74 .± 0.09 2.07 .± 0.17 0.12 .± 0.01 72.4 .± 0.8

Av45D4 30.0 .± 0.3 22.6 .± 0.7 0.05 .± 0.00 0.08 .± 0.00 107.7 ± 0.8 9.9 .± 0.1 16.5 .± 0.3 1.69 .± 0.07 2.91 .± 0.11 nd 24.9 .± 0.7

Av50D4 79.2 .± 1.1 121.3 .± 2.7 0.17 .± 0.01 0.23 .± 0.00 245.3 ± 2.6 30.8 .± 0.3 64.6 .± 0.7 838 .± 0.09 5.47 .± 0.24 0.25 .± 0.03 167.7 .± 2.3

Av61D4 39.3 + 0.4 15.0 + 0.5 0.05 + 0.01 0.14 + 0.00 117.0 + 1.0 8.9 + 0.0 19.1 + 0.2 1.47 + 0.04 4.12 + 0.12 0.08 + 0.00 31.4 + 0.6

@;for explanation of ~.ample number refer to APPENDIX Ia, Ib and Ic.

nd: not detected. Zn detection limit: 0.02 mg/lOOg.

... 8i

... a, a,

APPENDIX IVb: Selected elemental composition of termite mounds sampled at Daly River (sites: 1 & 3) and Howard Springs (site 6), following 0.1 N pepsin- IICI (pH 1.35)

• extraction on 2mm fraction size samples. Species: Tumulitermes pastinator.

Sample Element + sd (n-3) mg!lOOg

Number@ AI Ca Co Cu Fe K Mg Mn Na Zn Fe{II}

Tp02Dl 27.4 .± 0.2 17.9 .± 0.3 nd 0.04 .± 0.00 13.2 .± 0.2 11.4 .± 0.4 10.0 .± 0.1 1.19 .± 0.02 0.82 .± 0.13 0.09 .± 0.04 8.01 .± 0.26

Tp16Dt 25.4 .± 0.2 11.4 .± 0.1 nd 0.03 .± 0.00 9.8 .± 0.0 8.6 .±. 0.1 7.6 .± 0.2 1.24 ± 0.01 0.92 .± 0.09 0.01 .± 0.01 4.52 .± 0.05

Tp23DI 36.5 .±. 0.8 22.0 .±. 1.6 nd 0.05 ± 0.01 16.6 .± 0.5 12-9 .± 0.4 8.2 .± 0.3 1.18 .±. 0.05 3.80 .± 0.22 0.01 .± 0.02 9.72 ± 0.30

Tp34D3 23.7 .±. 0.2 25.0 .± 0.9 0.04 .± 0.01 1.45 .± 0.07 12.3 .± 0.2 20.9 .± 0.8 36.1 .± 0.3 2.36 .± 0.11 1.13 .± 0.05 0.76 .± O.o3 9.31 .± 0.26

Tp36D3 37.1 .± 1.4 15.1 .± 0.1 0.04 .± 0.01 0.95 .± 0.04 29.0 .± 1.3 14.9 .± 0.2 27.3 .± 0.6 1.94 .± 0.00 2.25 .± 0.35 0.82 .± 0.02 20.46 .± 1.32

Tp38D3 27.9 + 0.9 19.4 .± 0.2 0.04 .± 0.00 0.41 .± 0.04 12.6 .± 0.3 16.5 .± 0.8 2.'~ .± 0.4 2.87 .± 0.11 1.94 .± 0.()9 0.39 .± 0.02 8.83 .± 0.37

Tp45H 55.9 .± 0.2 22.1 .± 0.5 0.05 .± 0.00 0.23 .± 0.01 8.5 .± 0.2 4.0 .± 0.3 7.7 .± 0.2 2.65 .± 0.03 0.96 .± 0.20 0.06 .± 0.00 4.75 .± 0.29

Tp60H 39.0 .± 0.3 59.8 .± 1.8 nd 0.13 .± 0.01 10.2 .± 0.1 7.6 .± 0.4 17.9 .± 0.3 1.36 .± 0.02 1.27 .± 0.25 0.11 .± 0.01 9.63 .± 0.23

Tp63H 37.7 + 0.7 27.5 + 0.4 0.01 + 0.02 O.o? _+ 0.00 7.3 + 0.1 3.4 + 0.2 9.2 + 0.1 0.50 + 0.00 0.51 + 0.05 0.04 + 0.00 4.30 + O.o?

@; for e11:planation or sample number refer to APPENDIX Id, le and If.

nd: not detected. Co detection limit = 0.02 mgftOOg

APPENDIX lYe: Selected elemental composition of tennite mounds sampled at Daly River (3 & 4) and Howard Springs (site 6), following 0.1N pepsin- IICI (pH 1.35) extraction on 2mm fraction size samples. Spedes: Nasutitermes triodiae.

Sample

Number@

Element + sd (n=3) mg/100g

AI Ca Co Cu

25.8 .± 1.1 17.3 .± 0.4 0.01 .± 0.01 0.12 .± 0.01

29.9 .± 0.6

29.8 + 0.8

23.4 .± 0.6

18.7 .± 0.2

20.7 .± 0.2

29.6 + 0.4

47.4 .± 13

33.3 .± 0.1

0.03 .± 0.00 0.15 .± 0.01

0.02 .± 0.00 033 .± O.Q3

O.o2 .± 0.00 0.09 .± 0.00

o.ot .± o.oo o.o8 .± o.oo 25.1 .± 0.8 24.7 .± 0.3 0.03

20.8 .± 0.5 34.6 .± 0.8 0.04

.± 0.00

.± 0.01

nd

O.o7 .± 0.01

0.09

0.11

.± 0.00

.± 0.02 21.2 .± 1.0 32.2

20.8 .± 1.0 34.8

.± 0.9

.± 1.6 0.08 .± 0.00

12.7

15.8

225

Fe

.± 0.5 24.6

.±. 0.4 243

.± 0.2 25.2

21.7 .± 0.5 60.6

11.0 .± 0.2

12.1 .±. 0.2

30.4

12.9

20.1

623

50.7

K

.± 0.3

.± 03

± 0.2

.± 0.5

.± 0.6

.± 0.2

19.5 .± 0.2 24.6 + 0.0

0.03

0.04

0.03

.± 0.01

.± 0.01

.± 0.01

nd

0.12 .± 0.01 . 30.0

+ 0.6

.± 0.3

.± 0.9

.± 0.2

.± 0.5

.± 0.3

56.8 .± 1.1

51.2 .± 0.4

36.7 .± 0.7

31.7 .± 0.5

73.4

17.3

14.9

15.3

47.7

.± 0.9

.± 0.8

26.1

11.8 .± 0.8

.± 0.1

+ 0.2 16.7 + 0.4 0.03

.± 0.2

.±1.2 31.8

52.8

.± 0.6

.± 0.5

0.03 .± 0.00

.± 0.00

nd

0.12 .± 0.00

0.07

0.08

.± 0.01

.± 0.00

0.07 .±. 0.01

39.5

14.8

43.0

26.9 .± 03

.± 0.7

23.8 .± 0.1 27.8 .± 0.8

17.0 .± 0.2 45.6 + 0.4

9.8 .± 0.1

M_g_____ Mn Na Zn Fe(II)

39.3 .± 0.9 1.63 .±. 0.08 1.66 .± 0.03 0.10 .±. 0.01 10.92 .± 0.32

42.8 .± 0.4 2.22 .± 0.10 2.52 .± 0.28 0.13 .± 0.03 13.18 .± 0.40

88.2 .± 1.9 3.48 .± 0.11 6.30 .± 032 0.25 .± 0.01 18.32 .± 0.43

67.0 .± 1.1 3.29 .± 0.09 16.90 .± 0.37 0.05 .± 0.00 18.09 .± 0.24

42.0 .± 0.7 2.23 .± 0.01 7.68 .±. 0.17 0.09 .± 0.01 8.18 .± 0.20

14.3 .± 0.2 1.40 .± O.ot 3.05 .± 0.14 0.08 .±. 0.01 8.69 .± 0.18

27.1 .± 0.9 2.86 .± 0.03 3.77 .± 0.10 O.o? .± 0.00 23.61 .± 0.66

27.7 .± 0.7 2.11 .± 0.06 5.90 .± 0.25 0.08 .± 0.02 9.82 .± 0.20

32.8 .± 0.9 2.60 .± 0.11 2.36 .± 0.14 0.09 .± 0.00 16.18 .± 0.88

25.6 .± 0.2 2.80 .± 0.07 3.32 .± 0.06 0.14 .± 0.01 18.87 .± 0.16

26.9 .± 0.5 2.73 .± 0.07 1.72 .± 0.03 0.10 .± 0.01 24.75 .±. 0.23

30.2

31.3

50.0

13.3

.± 0.4

.± 0.6

1.15

1.28

.± 0.4 2.15

3.50

5.55

6.83

1.70

.± 0.23

.± 0.18

0.07 .±. 0.01

0.09 .± 0.06

.± 0.10 0.05 .± 0.01

.± 0.09

NtlOD3

Nt12D3

Nt14D3

Nt25D4

Nt26D4

Nt28D4

Nt31D4•

Nt34D4

Nt56D4

NlWD4•

Nt59D4•

Nt65D4

Nt68D4•

Nt72D4

Nt92H

Nt99H

Nt106H

44.4 .± 0.7 793 .± 1.2 nd

0.11

0.14

.± 0.01

.± 0.00

+ 0.00

14.5

16.6

13.8

+ 0.2

.±. 0.1 10.9 .± 0.2 213

.± 0.1

.± 0.2

1.10

1.81

.± 0.03

.± 0.05

.± 0.08

.± 0.03

.± 0.02 1.01 .± 0.03

0.18

0.26 .± 0.02

.±. 0.02

12.43

16.09

12.57

13.58

15.19

1234

.± 0.13

.± 0.15

.± 0.25

.± 0.17

.± 0.36

+ 1.28 53.0 + 0.7 91.7 + 1.9 0.08 + 0.02 0.15

@: [or explanation or sample number refer to APPENDIX Tg, Jh and li.

Underlined: Sample collected on the outside or the mound (0-lcm)

+ 0.3 16.4 + 0.2 23.6 + 0.5 3.35 + 0.09

•: Newly built mound material nd :not detected. Co detection limit =- 0.02 mgllOOg

2.01 + O.o7 0.23 + O.ot

... a, ....

.... ~

Appendix IVd: Selected elemental composition of termite mounds sampled at Daly River (site 1), following 0.1N pepsin-HCI (pll1.35) extraction

on 2mm fraction size samples. Species~ (Tumulitermes luutllls)


Number@ At Ca Co Cu Fe K Mg Mn Na Zn Fe(II)

Th01D1 34.71 .± 0.27 82.07 .± 0.84 O.o3 .± 0.01 O.Q7 .± 0.00 21.69 .± 0.67 25.44 .± 0.35 32.98 ± 0.33 4.34 .± 0.06 1.92 ± 0.12 0.10 .±. O.ot 19.62 .±. 0.46

Th02D1 35.74 ± 0.43 69.92 ± 2.12 0.04 .± 0.00 O.o7 .± 0.00 20.47 .± 0.40 10.91 ± 0.32 21.34 .± 0.83 3.03 .± 0.12 0.74 ± 0.04 0.16 .±. 0.08 17.38 .± 0.31

Th03Dl 30.13 .±. 033 29.42 .± 1. 15 nd 0.03 .± 0.00 14.36 .± 0.05 13.58 .± 0.11 13.67 ± 0.2.1 1.55 .± 0.05 1.01 ± 0.12 0.05 .± 0.01 1252 .± 0.57

@:for explanation of aample number refer to APPENDIX lj.

nd: not detected. Co detection limit: 0.02 mg/lOOg

APPENDIX IVe: Selected elemental composition of soils (0-10cm depth) sampled at Elliott (site 5), Daly River (sites: 1-4) and lloward Springs (site 6)

following 0.1N pepsin-HCI (pH 1-35) extraction on Zmm fraction size samples.

Sample Element + sd (n-3) mg!IOOg

Number AI ca Co Cu Fe K Mg Mn No Zn Fe{II)

OlE 9.50 .±. 0.32 45.70 .±. 1.26 nd O.o7 .±. 0.01 4.95 .±. 0.33 9.79 .±. 0.27 10.21 .±. 0.04 1.71 .±. 0.03 0.83 .±. 0.38 nd 0.76 .±. 0.09

02E 12.08 .±. 0.31 50.35 .±. 0.42 0.02 .±. 0.00 0.10 .±. 0.01 5.72 .±. 0.38 10.42 .±. 0.14 13JJ6 .±. 0.20 2.26 .±. 0.02 0.87 .±. 0.26 nd 0.91 .± 0.03

0501 26.28 .±. 0.54 2.78 .± 0.33 nd 0.01 .± 0.02 5.63 .± 0.46 2.35 .± 0.27 0.35 .± 0.01 0.46 .±. 036 0.55 .±. 0.45 0.10 .± 0.10 1.98 .± 0.20

0701 37.86 .±. 0.25 5.44 .±. 0.08 nd 0.01 .t. 0.01 7.45 .± 0.21 2.23 .± 0.19 0.69 .±. 0.04 0.27 ± 0.00 0.57 .± 0.08 0.02 .± 0.02 2.23 .±. O.o7

0902 49.00 .± 1.86 6.83 .± 0.15 0.04 .± 0.00 0.12 .± 0.01 34.82 .± 2.80 4.63 .;tO.o9 3.44 .±. 0.11 0.89 .± 0.03 0.26 .±. 0.04 0.03 .± 0.00 5.03 .±. 0.21

1002 51.51 .± 1.04 1.74 ± 0.08 0.03 .±. 0.01 0.09 .±. 0.01 16.33 .±. 0.51 2.91 .± 0.45 0.94 .± 0.03 0.31 .±. 0.01 1.28 .±. 0.02 0.03 .± 0.01 3.34 .± 0.04

1703 27.13 .± 0.56 9.30 .± 0.10 nd 0.05 .±. 0.01 13.91 .±. 0.72 9.35 .± 0.11 18.92 .± 0.03 0.94 .±. 0.04 1.77 .± 0.12 0.14 .± 0.03 7.21 .± 0.46

2203 35.24 .±. 0.16 6.56 .± 0.06 0.04 .± 0.01 0.11 .± 0.01 32.61 + 0.14 5.28 .t. 0.13 8.61 .±. 0.11 0.77 ± 0.01 1.17 .± 0.35 nd 4.99 .±. 0.51 2404 47.31 .±. 0.71 2.63 .± 0.05 0.02 .± 0.00 0.09 .± 0.01 33.56 .± 1.01 7.16 .±. 0.11 4.43 .±. 0.04 0.15 .±. 0.00 1.58 .±. 0.36 0.04 ± 0.00 7.63 .± 0.49

2604 26.59 .± 0.42 2.52 .± O.o? 0.02 .± 0.01 0.04 .±. 0.01 15.69 .±. 0.34 4.53 .±. 0.36 3.22 .± 0.05 0.13 .±. 0.01 1.35 .±. 0.32 0.03 .±. 0.02 5.33 ± 0.22 27H 46.37 .± 1.02 31.54 ± 0.54 nd 0.09 .±. 0.00 5.33 .±. 0.19 5.03 .± OJJ6 7.76 ± 0.12 1.17 .±. 0.05 1.08 .± 0.03 0.48 .±. 0.04 3.53 _±0.22

29H 69.26 + 2.18 43.63 + 2.97 0.06 + 0.01 0.10 + 0.00 7.16 + 0.30 1.90 + 0.06 5.75 + 0.31 2.17 + 0.09 1.21 + 0.14 1.91 + 0.06 4.19 + 0.26

For selected termite specie!! studied at each site, refer to APPENDIX lk. nd: not detected. Co and Zn detection limit • 0.02 mg/IOOg

.... ~

.... ... <:>

APPENDIX Va: Selected elemental composition of termite mounds sampled at Elliott (site 5), Daly River (sites: 2 & 4) following 0.1 N pepsln-JICI

(pll1.35) extraction and neutralisation (pH 7.50), puformed on 2mm fraction size samples.



Number AI Ca Co Cu Fe K Mg Mn Zn Fe(Il) A vOlE 0.70 .t 0.24 119.4 ± 0.9 0.05 .±. 0.01 0.07 .± Q.()() 0.39 ± 0.15 18.5 .±. 0.4 25.0 .± 0.7 4.00 ± 0.32 nd nd Avt5E 0.89 .±. 0.09 93.0 .±. 1.7 nd 0.06 .±. 0.01 0.10 ± 0.10 19.7 .±. 0.3 30.8 .± 0.9 1.36 ± 0.32 nd nd Av22E 1.68 ± 0.46 111.5 ± 0.2 0.03 .±. 0.00 0.07 .± 0.01 1.50 .± 0.75 13.9 .± 0.2 23.2 .± 1.3 2.51 ± 0.54 nd 2.01 .±. 0.27 Av3102 0.37 ± 0.17 27.1 ±1.1 0.03 ± 0.01 0.05 .±. 0.02 0.73 ± 0.06 14.0 .± 0.1 11.5 ± 0.4 2.65 ± 0.37 nd 0.84 ± 0.10 Av34D2 0.94 ± 0.57 12.5 ±0.8 nd nd 0.35 ± 0.16 5.9 ± 0.2 4.6 .± 0.1 0.80 ± 0.31 nd 1.31 ± 0.29 Av39D2 0.94 ± 0.53 20.3 .±. 1.6 nd 0.04 .±. 0.00 1.64 .±. 0.27 7.3 .± 1.0 7.7 .± 0.4 221 ± 0.55 nd nd Av4504 1.19 .± 0.29 17.5 ± 0.2 nd nd 3.68 ± 1.36 9.6 ± 1.8 14.4 ± 0.1 0.78 ± 0.06 nd 0.40 .± 0.01 Av5004 231 .± 0.38 87.4 .± 26 0.04 .± 0.00 0.07 .± 0.01 8.99 .± 286 29.0 .± 0.4 54.0 .± 1.1 3.20 .± 0.50 nd nd Av61D4 0.44 + 0.16 11.9 + 0.2 nd 0.04 _± 0.01 0.77 _± 0.31 7.8 _± 0.2 16.5 + 0.2 0.54 +_ 0.03 nd nd

@:for explanation of sample refer to Appendix Ia, lb and !c.

nd: not detected. Detection limit (mg/lOOg): A1 = 0.05; Co and Zn = 0.02

Cu = 0.01; Fe .. 0.06; Fe(li) = 0.2.

APPENDIX Vb• Selected elemental composition of termite mounds sampled at Daly River (1 & 3) and lloward Springs (site 6), following

O.LN pepsin-IICI (pH 1.35) extraction and neutralisation (pll 7.50) on 2mm fraction size samples.

Species: Tumulilermes pastinalor.

Sample Element + sd (n-3) mg/100g

Number@ AI Ca Co Cu Fe K M~ Mn Zn Tp02D1 0.56 ..±. 0.2 17.3 .± 0.3 nd 0.01 .± 0.01 0.04 .± 0.1 10.7 .± 0.4 9.3 .± 0.2 0.83 .± 0.03 nd Tp16Dl 0.88 .± 0.3 10.8 .± 0.2 nd nd 0.11 .± 0.1 8.3 .± 0.3 7.0 .± 0.1 0.80 + 0.05 nd Tp23Dl 0.70 .±. 0.2 19.8 .± 1.0 nd 0.01 .± 0.02 0.23 .±. 0.1 12.4 .±. 0.5 7.3 .± 0.2 0.74 .± 0.01 nd Tp34D3 0.74 ± 0.3 18.8 .± 0.3 nd 0.10 .±. 0.00 nd 17.3 .± 0.5 22.4 .±. 0.4 2.32 .±. 0.09 nd Tp36D3 0.65 .±. 0.2 24.6 .±. 0.8 nd 0.21 .± 0.04 nd 14.9 .± 0.3 33.6 .±. 1.8 1.74 .± 0.47 nd Tp38D3 1.57 ± 1.7 14.4 .± 0.4 nd 0.13 .± 0.01 nd 15.5 .± 0.2 25.1 .± 1.9 1.27 .±. 0.35 nd Tp45H 1.12 .±. 1.4 19.9 .± 0.6 nd 0.03 .± 0.00 nd 3.5 .± 0.4 7.1 .±. 0.1 1.77 .± 0.23 nd Tp60H 1.22 .±. 0.5 57.4 .± 1.8 nd 0.05 .± 0.00 0.09 .± 0.0 7.1 .± 0.9 17.9 .±. 0.4 0.95 .± 0.05 nd Tp63H 0.54 + 0.3 27.9 + 0.7 nd nd nd 20 + 0.1 9.0 + 0.2 0.36 + 0.05 nd

@; for explanation of sample refer to Appendix Id, Ie and Ir.

nd: not detected. Detection limit (mg/lOOg); Co and Zn = 0.02; Fe(II) .. 0.2

Fe(ll) nd nd nd nd nd nd nd nd nd

"' .... ~

APPENDIX Vc: Selected elemental composition of termite mounds sampled at Daly River (sites: 3 & 4) and Howard Springs (site 6), following 0.1N pepsfn-IICI ... extraction and neutralisation (pll 7.50) on 2mm fraction size samples. Species: Nasutitermes triodiae. tj

Sample

Number@

Element + sd (n=3) mg1100g

Nt!OD3 Nt12D3 Nt1403

AI Ca

0.55 ± 0.12 16.8 .± 0.4 .± 0.3 0.67

0.79 .± 0.06 .± 0.26

19.4 27.5 ± 0.6 0-03

Co

nd nd .± 0.01

0.01 0-03 0.11

Cu

.± 0.02

.± 0.03

.± 0.01 " Nt25D4

Nt26D4 Nt2804 Nt31D4' Nt34D4 Nt56D4 Nt59D4' Nt65D4

10.79 .± 2.84 42.7 .± 0.3 0.03 .± 0.00 0.07 .± 0.00 2.57 .± 0.20 26.9 .± 0.0 0.34 .± 0.32 21.1 .± 0.6 3.44 .± 2.43 29.7 .± 0.6 3.10 .± 0.64 31.6 .± 0.6 1.39 .± 0.33 30.2 .± 1.4 0. 75 .± 0.38 21.4 .± 0.3 1.80 .± 1.50 15.8 + 0.9

nd nd nd nd nd nd nd

_Nt68D4' 0.90 .± 0.20 14.7 .± 0.4 nd Nt72D4 4.64 .± 0.18 30.4 .± 0.9 0.02 .± 0.02 Nt92H 1.37 .± 0.85 50.4 .± 1.9 nd Nt99H 231 .± 0.30 78.5 .± 2.4 nd Nt106H 0.90 + 0.18 80.2 + 1.2 0.02 + 0.01 @ : [or explanation or sample re[er to Appendix lg, lh and Ii

Underlined: sample collected on the outside o( the mound (0-lcm)

• : Newly built mound material .

0.03 .± 0.00 0.03 .± 0.01 0.05 .± 0.00 0.05 .± 0.01 0.03 .± 0.00 0.04 .± 0.00 0.02 .± 0.01 0.03 .± 0.00 0.05 .± 0.01 0.03 .± 0.00 0.06 .± 0.00 0.05 + 0.00

Fe K Mg Mn Zn 0.16 .± 0.06 13.3 34.6 .± 2. 7 0.97 + 0.33 0.12 .± 0.04 13.6

.± 0.2

.± 0.8 37.3

85.7 .± 3.5 .± 2.2

1.38

2.90 .± 0.48 .± 0.09

nd nd nd nd nd

0.79 9.56 1.45

4.48

.± 0.39 23.1 .± 0.5 + 0.59 .± 0.16 nd .± 2.14

57.1 63.1

.± 0.4 60.0 .± 3.3 2.40 .± 0.49

.± 28 34.7 .± 0.6 1.77 .± 0.05 50.1 .± 0.7 11.8 .± 1.2 0.78 .± 0.26 0.04 58.1 .± 1.2 24.2 .± 0.5 1.90

.± 0.04

t.99 .± 0.22 0.48 .± 0.45

50.7 36.8

.± 1.4 27.7 .± 1.3 1.70

.± 0.7 28.4 .± 1.3 .± 0.3

1.28 1.68

.± 0.06

.± 0.05

.± 0.36

.± 0.24

0.03 .± 0.03 nd nd

31.7 .± 0.5 22.9 nd 0.81 0.83

.± 0.04

.± 0.21 27.2 .± 0.2 26.3 .± 1.5 0.68 .± 0.32 0.04 .± 0.01 1.44 .± 0.19 27.8 .± 1.6 28.0 .± 0.8 4.80 .± 0.53 44.7 .± 0.5 48.5 .± 1.3 0.20 .± 0.05 9.6 .± 1.2 12.6 .± 0.6 0.82 .± 0.06 10.6

nd 16.1 ~ : n=Z

+ 0.4 + 114

20.4 2113

.± 0.4 + 113

0.79 .± 0.10 0.01 .± 0.03 1.76 .± 0.06 nd 0.68 .± 0.13 nd 1.26 .± 0.08 nd 213 + 0.05 nd

nd: not detected. Detection limit (mgflOOg): Q, and Zn = 0.02; Cu = 0.01;

Fef!J) nd nd

0.32 .± 0.11 nd

0.41 .± 0.05 nd

0.61 .± 0.12 0.45 .± 0.02 0.41 .± 0.02 0.40 ± 0.10 0.37 .± 0.03 0.44 .± 0.01 0.45 ± 0.03

nd 0.24 ± 0.02

nd

Appendix Vd: Selected elemental composition or termite mounds sampled at Daly River (site 1) (Tumulitermes hastilis)

rollowing 0.1N pepsin-HCI {pH 1.35) extraction and neutralisation (pH 7.50) on 2mm rractJon size samples.

Sample

Number@ AI Ca Co

ThOID1 275 ± 0.31 71.65 ± 2.19 nd

Th02D1 3.56 ± 2.25 61.n ± 2.03 nd

Th03D1 2.35 .± 1.57 26.00 .± 0.63 nd

@: (or explanation of sample number refer to APPENDIX Jj.

nd: not detected. Co and Zn detection limit: 0.02 mg/lOOg

Element +

Cu

0.01 ± 0.01

0.02 .± 0.02

0.02 .± 0.02

sd (n=3) mg/100g

Fe K Mg Mn

2.61 .± 0.11 25.75 .± 2.80 30.26 ± 1.01 3.02 .± 0.17

2.33 .± 0.73 10.02 .± 0.40 18.89 .± 0.44 209 ± 0.14

1.26 .± 0.36 12.81 .± 0.69 11.88 .± 0.06 1.00 ± 0.04

Zn FeQI)

nd 0.55 .± 0.04

nd 0.55 .± 0.10

nd 0.52 .± 0.06

... <;!

.... ~

APPENDIX Ve: Selected elemental composition of soils (0-lOcm depth) sampled at Elliott (site 5), Daly River (sites: 1-4) and Jloward Springs (site 6),

following O.lNpepsin-IICI (pll1.35) extraction and neutralisation (pll 7.50) on 2mm fraction size samples.

Sample Element + sd (n~3) mg/!OOg

Number AI Ca Co Cu Fe K Mg Mn Zn Fe(II) OlE 0.60 .±. 0.14 42.29 .± 1.57 nd nd nd 1.13 .±. 0.21 9.38 .± 0.24 1.23 .± 0.02 nd nd 02E 0.62 .± 0.17 45.65 .±. 1.19 nd nd nd 7.80 .±. 0.95 11.75 .±. 0.37 1.68 .±. 0.11 nd nd 0501 0.52 .± 0.07 2.83 .± 0.03 nd nd nd nd 0.56 .±. 0.01 0.19 .±. 0.02 nd nd 0701 0.73 .± 0.11 5.51 .±. 0.32 nd nd nd nd 0.79 .±. 0.11 0.12 .±. 0.04 nd nd 09D2 0.05 .±. 0.09 5.65 + 0.14 nd nd nd 5.40 .±. 0.70 2.45 .±. 0.13 0.40 .± 0.12 nd · nd 10D2 0.86 .±. 0.21 1.54 .± 0.04 0.02 .±. 0.00 nd nd 0.72 .± 0.62 0.48 .±. 0.01 0.06 .±. 0.01 nd nd 17D3 0.26 .± 0.01 7.86 .±. 0.02 nd 0.01 .±. 0.01 nd 8.49 .±. 0.47 16.14 ± 0.24 0.63 .±. 0.02 nd nd 22D3 0.28 .±. 0.07 5.57 .±. 0.11 nd nd nd 4.18 .±. 0.30 7.17 .±. 0.19 0.41 .±. 0.04 nd nd 24D4 nd 2.27 .± 0.08 nd nd nd 8.50 .±. 209 3.55 .± 0.10 0.03 .± 0.01 nd nd 26D4 nd 1.91 .± 0.11 nd nd nd 6.96 .±. 2.88 2.46 .±. 0.07 O.D3 .±. 0.01 nd nd 27H 0.92 .±. 0.22 27.81 .±. 1.77 nd nd nd 3.23 .±. 0.61 6.26 .±. 0.85 0.64 .±. 0.15 nd nd 29H 0.91 + 0.09 39.16 + 3.08 0.03 + 0.01 nd nd 1.72 + 0.95 4.67 + 0.49 1.37 + 0.15 nd nd

For selected tennite spedes studied at each site, refer to APPENDIX lk..

nd: not detected. Detection limit (mgllOOg): AI and K = 0.05; Co and Zn = 0.02

Cu = 0.01; Fe = 0.06; Fe(II) = 0.2

"----... .----~ ----.

-"' 0 0 ~

' "' E ~

<(

-"' 0 0 ~

' "' E ~

"' 0

-"' 0 0 ~

' "' E ~

0 0

Pepsin-HCI (pH 1.35)

75 Site 1 Site2 Site3

60

45

30

15

0 Tp Th s Av s Tp Nt s Av Nt s Av s Tp Nt s

150 Site 1 Site2 Site3 Site4 SiteS Site6

100

50

0 Tp Th s M s Tp Nt s Av Nt s Av s Tp Nt s

0.15 Site 1 Site2 Site3 Site4 SiteS Site6

0.10

0.05

0.00 -'--,-~ Th S ~ S ~ ~ S ~ Nt S ~ S ~ ~ S

SPECIES I SOIL

275

APPENDIX VI-A Graphic representations of the soil - mound effects (mean ± SE) on element concentrations (mgllOOg) following the in vitro test on termitaria of Amitermes vitiosus (Av), Tumub1ermes pastinator (Tp), Tumulitermes hastilis (Th), Nasutitermes triodiae (Nt), and soil (S) (S) (0-IOcm), sampled at sites 1-6, following pepsin-HCI (pH 1.35) extractions: aluminium, calcium and cobalt.

276

-"' 0

Pepsin-HCI (pH 1.35)

1.30 -,-----,----,---,---,.----,------, Site 1 Site2 Site3 Slte4 Site 5 SiteS

1.04

0 0.78 ~

d, 5 0.52 ::> 0

-"' 0

0.26

0.00 I "i" C(l ..,.. I ~ Th S ~ S ~ M S ~ M S ~ S ~ M S

225-,-----,---,---,----,---,---~ Site 1 Site2 Site3

180

0 135 ~

E "' lL

g 0 ~

' "' E ~

=

90

45

0 I B!lll ~ Th S ~ S ~ M S ~ M S ~ S ~ M S

100 Sb6 Site 5 Site 3 Site 4 Site 1 Site 2

80

60

40

;f 20

0 I 171 171 "? I 171 "7' I ~ Th S ~ S ~ M S ~ Nl S ~ S ~ M S

APPENDIX VI-B

SPECIES I SOIL

Graphic representations of the soil • mound effects (mean ± SE) on element concentrations (mg/IOOg) following the in vitro test on tennitaria of Amitermes vitiosus (Av), Tumulitermes pastinator (Tp), Tumulitermes hastilis (Th}, Nasutitermes triodiae (Nt), and soil (S) (0-IOcm), sampled at sites 1-6, following pepsin-HCI (pH 1.35) extractions: copper, iron and iron (ii).

APPENDIX VI-C

SPECIES I SOIL

Graphic representations of the soil • mound effects (mean ± SE) on element concentrations (mgllOOg) following the in vitro test on tennitaria of Amitermes vitiosus (Av), Tumulitermes pastinator (Tp), Tumu/itermes hastilis (Th). Nasutitermes triodiae (Nt), and soil (S) (0-lOcm), sampled at sites 1-6, following pepsin-HCI (pH 1.35) extractions: potassium, magnesium and manganese.

278

12 Site 1

10

-CD 8 0 0 ~

' 6 CD E -"' 4 z

2

0

1.75 Site 1

I 1.40 -CD

0 0 ~

1.05

' CD E 0.70 -c N

0.35

0.00

APPENDIX VI- D

Pepsin-HCI (pH 1 .35)

I Site 2,/ Site3

I Site2j Site3 I Site4 j SiteS j SiteS

SPECIES I SOIL

Graphic representations of the soil - mound effects (mean ± SE) on element concentrations (mg!IOOg) following the in vitro test on tennitaria of Amitermes vitiosus (Av), Tumulitermes pastinator (Tp), Tumulitermes hastilis {Th), Nasutitermes triodiae (Nt), and soil (S) (0-lOcm), sampled at sites 1-6, following pepsin-HCI (pH 1.35) extractions: sodium and zinc

0

APPENDIX VII-A

Nt S Av S Tp Nt S

Site 5 Site 6

Graphic representations of the soil ~ mound effects (mean ± SE) on element concentrations (mg/1 OOg) following the in · vitro test on termitaria of Amitermes vitiosus (Av),Tumu/itermes pastinator {Tp), Tumu/itermes hastilis (Th), Nasutitermes triodiae (Nt} and soil (S) (0-!0m),sampled at sites l-6, following pepsin-HCI (phl.35) extractions and neutralisation (pH 7 .5): aluminium, calcium and cobalt.

280 pH 7.5 Filtrate

0.20 -,----,.---,------.----c--r----,

c;, 0.15 0 0 ~

' ~ 010 -::> 0 0.05

Q.QQ I ep

Site 4 SiteS Site 6

Av Nt S Av S Tp Nt S

7.50 -,----,-----,----r--,---,-------,-------, Site 1 Site 2 Site 3 Site 6

c;, 0

6.00

0 4.50 ~

do s 300

" u_

1.50

000 I '1" Tp Th S Av S Tp Nt S s

0.7 B I Sitol I Site2 I Sit• 3 I h•I• I <a. • I oa. < I - 060 OJ 0 0 :::: 0.45

]' 030 -

" u_ 0.15

1;j;1 I I 1;j;1 I tj'l tj'l I 0.00 I 1 I I I I I I

APPENDIX Vll-B

~ S ~ M S ~ M S ~ S ~ M S

SPECIES I SOIL

Graphic representations of the soil - mound effects (mean ± SE) on element concentrations (mg/1 OOg) following the in vitro test on termitaria of Amitermes vitiosus (Av),Tumulitermes pastinator (Tp), Tumulitermes hastilis (Th), Nasutitermes triodiae (Nt), and soil (S) (0-l Om),sampled at sites I-6, following pepsin-HC (ph 1.35) extractions and neutralisation (pH 7 .5): copper, iron and iron (Ii).

60 Site 1

50 ~ 40 a> 0 0 30 ~

' a> 20 E -

" 10

0

-10 Tp Th

0

APPENDIX Vli-C

pH 7.5 Filtrate 281

Sitel Site 3 Site 5 Site 6

Av s Tp Nt s Av Nt

s

SPECIES I SOIL

Graphic representations of the soil - mound effects (mean ± SE) on element concentrations (mg/1 OOg) following the in vitro test on termitaria of Amitermes vitiosus (Av),Tumu/itermes pastinator (Tp), Tumulitermes hastilis (Th), Nasutitermes triodiae (Nt), and soil (S) (0-lOm),sampled at sites 1-6; following pepsin-HCI (phl.35) extractions and neutralisation (pH 7.5): potassium, magnesium and manganese.

282

pH 7.5 Filtrate

O.D1 Site 1 Site 2 Site 3 Site 5 Site6

0.01 -Ol 0 0 001 ~ . a, 5 000 c N 000

000 ~ Th S ~ S ~ M S ~ M S ~ S ~ M S

SPECIES I SOIL

APPENDIX VII-D Graphic representations of the soil - mound effects (mean ± SE) on element concentrations (mg/IOOg) following the in vitro test on termitaria of Amitermes vitiosus (Av),Tumulitermes pastinator (Tp), Tumu/itermes hasti/is (Th), Nasutitermes triodiae (Nt), and soil (S) (0-IOm), -sampled at sites 1-6, following pepsin-HCI (phl.35) extractions and neutralisation (pH 7.5): zinc

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