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Southern Cross University

ePublications@SCU
Theses

2010

Phytochemistry, allelopathy and the capability


attributes of camphor laurel (Cinnamomum
camphora (L.) Ness & Eberm.)
John Robert Schenk
Southern Cross University, j.schenk.10@scu.edu.au

Suggested Citation
Schenk, JR 2010, 'Phytochemistry, allelopathy and the capability attributes of camphor laurel (Cinnamomum camphora (L.) Ness &
Eberm.)', PhD thesis, Southern Cross University, Lismore, NSW.
Copyright JR Schenk 2010

ePublications@SCU is an electronic repository administered by Southern Cross University Library. Its goal is to capture and preserve the intellectual
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Phytochemistry, Allelopathy and the Capability


Attributes of Camphor Laurel (Cinnamomum
camphora (L.) Nees & Eberm.)
in north-eastern New South Wales.

By John Robert Schenk


B.A. (Hons.) (NSW.)

A thesis submitted for the degree of Doctor of Philosophy:


School of Environmental Science and Management
Southern Cross University

September 2009

ACKNOWLEDGEMENTS
There is much appreciation I wish to express to my supervisors Prof. Peter Saenger
(SCU) and Dr. Brett Stubbs (SCU) for their encouragement, guidance and
professional criticism during this research.

Special thanks should also be given to Dr. Lyndon Brooks (SCU) and Dr. Antony
McCardell (SCU) for guidance with the statistical methodology particularly the nonlinear regressions and deviance test performed on the vascular plant germination and
algal growth data. Dr Don Brushett (SCU) also needs special recognition for the work
undertaken on the gas chromatography and interpretation of results.

Much insight was gained from accessing research papers written in other languages
and I must thank Meg OReilly for the translation of the Russian paper, Tazuko
Mclaren for the Japanese papers, and the many Chinese post-graduate students in
Phytochemistry (SCU) who enabled access to documents written in Mandarin. The
assistance and friendship of the staff, technical officers, lecturers and fellow students
need also be recognized as they helped keep me happily working away, and listened
to my regular babble about the research.

My family should also be given a big hug for their understanding and assistance
during this work as I could not have undertaken it without this special help.

DECLARATION
I certify that the substance of this thesis has not already been submitted for any degree and is not
currently being submitted for any other degree or qualification, either in whole or in part. I certify that
any assistance received in preparing this thesis, and all sources used, have been acknowledged within
the content of the document.
I acknowledge that I have read and understood the Universitys rules, requirements, procedures and
policy relating to higher degrees research award and to my thesis. I certify that I have complied with
the rules, requirements, procedures and policy of the University.
Signed,

John Robert Schenk.


September 2009

ii

This thesis is dedicated to:

my mother Gwen, my son Justin, my partner Ella, the many people


who have provided inspiration and guidance through the years
and to the natural environment I dearly love.

iii

Abstract
The camphor laurel tree (Cinnamomum camphora (L.) Nees & Eberm.) was tested for
allelopathic influence in studies of seed germination, seedling growth and soil algal
populations of species identified in the regeneration assemblages of the Big Scrub
region in north-eastern New South Wales. This included an allelopathy glasshouse
trial on germinating seed and soil algae, the development of a technique for field
identification of the camphor laurel chemo-types, an application of the chemo-type
differentiation technique in a field assessment of allelopathic influence on seedling
growth, and a review of the trees capability attributes providing a greater
understanding of its role in plant succession.

The allelopathy glasshouse trial was performed on germinating seed from 52 vascular
plant species and 27 soil algal species associated with the camphor laurel
assemblages. Aqueous solutions obtained from macerated camphor laurel leaves of
the cineole and camphor chemo-types were prepared including nutrient and acidity
adjustments. These treatments were applied to the germinating seed until germination
was complete. Measurements of germination number and percentage over time and
growth measurements at completion of germination for radicle length, shoot length,
leaf number and leaf area were performed. Germination over time was modeled for
each species with a non-linear regression using the Richards function which provided
an asymptote, mid-inflection point, days to mid-inflection and germination rate. A
deviance test and residual analysis were undertaken for each of these modeled
variables which identified significant effects due to the extracts. In the statistical
assessment of growth measurements this used t-tests for each species and growth
attribute including a summary effect table which also identified significant effect.

For the field identification of the chemo-types several methods were used to
investigate possible differences including sectioning of the lamina and use of 11
histological stains, leaf chlorophyll extraction and precipitate comparison, leaf
venation and sclerophylly comparison and olfactory recognition of the chemo-types
using gas chromatography of the leaves compared with a blind olfactory test.

iv

In the field recognition of allelopathy, quadrating of seedlings found across various


sites where both chemo-types of camphor laurel and other vegetation types were
located produced measurements for shoot length, leaf number and leaf area.
Measurements were compared for each species and growth variable across these sites
using t-tests which provided a measure of significant influence due to the camphor
laurel canopy types on seedling growth.

The glasshouse trial identified that direct allelopathy originating from camphor laurel
leaves influences vascular plant seedlings through a significant delay to germination
and the significant reduction of radicle length, shoot length, leaf area and leaf number
in most native and exotic vascular species associated with camphor laurel plant
communities. No significant difference between the effects of the two chemo-types
was found. Alpha selection of plant species was indicated to be occurring through this
process of inhibition with potential to alter plant successional sequence in the field.
Soil algal population and growth in the glasshouse were also significantly influenced
by the allelochemics with many species either disappearing from the soil or
possessing reduced vigour. This was identified as being important, as soil algae assist
in many ecological functions such as soil wet-ability, moisture retention, humification
processes, nutrient fixation and release, seed germination enhancement and as a food
source for invertebrates. This indicates that allelopathy may also be indirect in its
operation on vascular plants through influencing soil algal succession which in turn
influences seed germination and plant growth, suggesting that Beta selection is also
occurring in the plant succession.

The testing for the field identification of the chemo-types resulted in olfactory
recognition being the most reliable, repeatable and inexpensive technique to assess
large numbers of camphor laurel trees, and was further applied in the field assessment
of allelopathy. During the process of gas chromatography 19 new oils were identified
in the leaves of camphor laurel.

The field assessment for allelopathy of seedling shoot length, leaf numbers and leaf
areas across the sampled sites found that significant reductions in all these growth
attributes occurs in many of the species including camphor laurel itself. This was
further verified against the glasshouse growth trial for most species indicating the
v

allelopathic effect is active but also appeared to be more obvious than in the
glasshouse trial. The comparison of effect between the chemo-types on field seedling
growth was only possible for camphor laurel seedlings, as the cineole chemo-type
was found to be confined to sporadic locations along creek lines which prevented
adequate sampling. It was found that growth in camphor laurel seedlings is
significantly reduced below a camphor laurel canopy of both chemo-types indicating
that greater competitive advantage for the species occurs outside the area of
allelopathic influence. The factor of rarity of cineole also suggested that the
camphor chemo-type was more invasive, being in greater abundance in the
regeneration assemblage and able to exist in drier ridgeline environments.

An extensive number of capability attributes was identified for camphor laurel thereby
indicating that the tree is both highly stress tolerant and highly competitive,
contributing to the influence of allelopathy and invasiveness of the tree. The process
of allelopathy was seen to provide ecological advantage to the camphor laurel tree by
reducing the competitiveness of the surrounding vegetation which provides a
facilitated growth of the species, as well as a factor contributing to the trees
dominance and persistence.

Ultimately, the findings indicate that camphor laurel does have a significant influence
in altering plant succession. Therefore as the species expands its presence in the
region, habitat change will also increase, placing further pressure or threat on both
plant and animal communities. The work also suggests that there is a need to seek a
biological control, assess the trees impact on leaf litter invertebrates and water
quality, and to improve management of camphor laurel recruitment in forested areas.

vi

Phytochemistry, Allelopathy and the Capability


Attributes of Camphor Laurel (Cinnamomum
camphora (L.) Nees & Eberm.)
in north-eastern New South Wales.
TABLE OF CONTENTS
VOLUME I

Page

TITLE PAGE

ACKNOWLEDGEMENTS

ii

DECLARATION

ii

DEDICATION

iii

ABSTRACT

iv

LIST OF TABLES

xi

LIST OF FIGURES

xii

LIST OF PLATES

xv

LIST OF APPENDICES

xv

LIST OF ABBREVIATIONS

xvi

CHAPTER 1 - INTRODUCTION

1.1

Study Background

1.2

Capability Attributes of Camphor Laurel

1.2.1 Brief Physical Description and Recent Classifications


1.2.2 Geographical Range and Climate
1.2.3 Ecological Strategies of Camphor Laurel
1.2.4 Summary of Ecological Characteristics

9
11
15
41

vii

1.3

Further Information Needs

43

1.3.1 Introduction
1.3.2 Definition of Allelopathy
1.3.3 Allelopathy and Camphor Laurel
1.3.4 Research Hypothesis and Approach
1.3.5 Significance of the Research

43
44
47
54
56

CHAPTER 2 - THE INFLUENCE OF AQUEOUS LEAF


EXTRACTS OF CAMPHOR LAUREL ON THE
GERMINATION OF SEED AND GROWTH OF SOIL
ALGAE

57

2.1

Introduction

57

2.2

Aim and Hypothesis

58

2.3

Methods

58

2.3.1 Experimental Design


2.3.2 Experimental Unit and Volatile Loss
2.3.3 Environmental Parameters
2.3.4 Extract Preparation and Application
2.3.5 Vascular Plant Selection, Treatment and Planting
2.3.6 Algal Selection, Inoculation and Treatment
2.3.7 Data Types Collected
2.3.8 Statistical Approach
2.3.9 Vermin Control and Fungal Invasion

58
59
61
62
64
65
66
67
70

Results

71

2.4.1
2.4.2
2.4.3
2.4.4
2.4.5

71
71
74
74

2.4

2.4.6
2.4.7

2.5

2.6

Propagation Unit Temperature


Chemistry of the Vat Solutions
Vascular Species Trialed
Germination - Pre-statistical Results
Soil Algae Trialed and Pre-statistical Results of
Growth
Seed Germination and Algal Growth- Significant
Effects of the Treatments
Vascular Plant Growth - Significant Effects of the
Treatments

97
104
107

Discussion

120

2.5.1
2.5.2
2.5.3

120
124
127

Vascular Plants
Soil Algae
Environment Control and Experimental Limitations

Conclusion

129
viii

CHAPTER 3 - FIELD RECOGNITION OF THE


CAMPHOR LAUREL CHEMO-TYPES

132

3.1

Introduction

132

3.2

Aim

133

3.3

Methods

133

3.3.1
3.3.2
3.3.3
3.3.4
3.3.5

133
136
137
137
137

3.4

Sectioning and Staining


Leaf Chlorophyll Extraction
Venation
Leaf Sclerophylly
Olfactory Recognition

Results

138

3.4.1
3.4.2
3.4.3
3.4.4
3.4.5

138
139
143
143
143

Sectioning of Lamina
Leaf Chlorophyll Extraction
Venation
Leaf Sclerophylly
Olfactory Recognition

3.5

Discussion

144

3.5

Conclusion

147

CHAPTER 4 - FIELD ASSESSMENT OF CAMPHOR


LAUREL ALLELOPATHY IN VASCULAR
PLANT SEEDLINGS

149

4.1

Introduction

149

4.2

Aim and Hypothesis

150

4.3

Methods

151

4.4

Results

153

4.4.1
4.4.2
4.4.3

154
154
163

Sites, Quadrats and Sample Size


Seedling Measurements
Significant Effects below the Canopy

4.5

Discussion

165

4.6

Conclusion

169

ix

CHAPTER 5 CAMPHOR LAUREL: POTENTIAL AND


POSSIBILITIES

171

5.1

Outcomes of the Phytochemistry and Allelopathy Research

172

5.2

The Combined Capability Attributes of Camphor Laurel


An Increasing Threat to Plant and Animal Biodiversity

172

Management and Further Research

174

5.3

REFERENCES

179

LIST OF TABLES
Table No.
1.

2.

3.

4.

5.

6.

7.

Title

Page

Temperature and precipitation for the provinces of southern


China in which camphor laurel is naturally distributed
according to Chen et al. (2004).

14

Temperature and precipitation for four Australian


locations where weed invasion resulting from camphor
laurel has been recorded by Firth (1979) and Batianoff and
Butler (2002).

14

Comparison of frugivores identified as present in the


camphor laurel forests by Nielan (2004) using the feeding
guild structure applied in that study, compared to known
dispersers of camphor laurel seed cited in the literature.

36

Summary of the competitive and stress-tolerant attributes


applied to camphor laurel following Grimes C-S-R strategy
(1979).

42

Confidence Interval at 95% (0.05) and Standard Deviation of


the oils of camphor laurel in: (a) cineole chemo-type;
(b) camphor chemo-type.

51

Vascular plants trialed and their seed characteristics


arranged by seed weight.

75

Percentage germination of the trees and shrubs in the


various treatment response groups.

77

Percentage germination of the vines and herbs in the


various treatment response groups.

78

The distinct germination response groups, species (a) and


seed characteristics (b) influenced by the cineole and
camphor treatments.

94

10.

Soil algae identified in the first algal trial.

99

11.

The most influenced vascular species by the treatments in the


time taken to reach mid-inflection.

121

Histological stains, substances stained and references to


staining techniques used in the application of the systematic
sectioning and staining of camphor laurel leaves.

135

8.

9.

12.

xi

13.

14.

15.

Comparison of secretory cells and oil globules of the


leaf in two gas chromatographed chemo-types of camphor
laurel using Sudan III stain.

141

Comparison of the oil globules in four trees of camphor laurel


whose chemo-types were assessed using Sudan III staining,
followed by verification through blind testing against gas
chromatography.

141

The identification of the camphor laurel chemo-types


using olfactory recognition during blind testing of gas
chromatographed leaves.

144

LIST OF FIGURES
Figure No.
1.

Title

Page

Biosynthetic pathways identified as probable origins of


allelochemic agents.

46

Camphor laurel leaf oil (terpenoid) content of the: (a) cineole;


(b) camphor chemo-types.

50

Quantitative comparison of the phytotoxic phenolics


from the leaf of *Cinnamomum camphora.

52

Propagation unit used for the germination of seed and growth


of algae in the allelopathy glasshouse trial.

60

5.

Propagation unit temperature on a daily and seasonal basis.

71

6.

Total nitrogen (a); total phosphorus (b) of the vat solutions


prepared for the control, cineole and camphor treatments
applied to seeds.

72

2.

3.

4.

7.

8.

9.

pH adjustments of the four vat solutions prepared for the


treatments applied to seeds for: (a) Vat solution 1; (b) Vat
solution 2; (c) Vat solution 3; (d) Vat solution 4.

74

Comparison of the highest overall germination response in


the treatments for each species.

79

Number and percentage of variability in: (a) commencement


of germination and (b) completion of germination compared to
the control.

96

xii

10.

Number and percentage of species in various response groups


compared against the control for the cineole and camphor
treatments.

97

11.

Number of plant families in each response group.

97

12.

Growth of algae over the media surface expressed as percentage


cover over time for: (a) sample 1; (b) sample 2; (c) sample 3.

100

Diversity and mean abundance of algae across the treatments


in algae sample 1.

101

Sample 1 algae of high relative abundance in the control and


decreased relative abundance in the treatments.

102

Sample 1 algae of high relative abundance in the control and


differential abundance in the treatments.

102

Sample 1 algae of low relative abundance in the control and


absent in the treatments.

103

Sample 1 algae either absent or with low relative


abundance in the control and increased relative
abundance in the treatments.

103

Sample 1 algae of low relative abundance in the control and


differential abundance in the treatments.

103

Total number of tree, shrub, vine and herb species showing


significant variation in the post-emergent measurements compared
to the control for shoot length (a); radicle length (b); leaf
number (c); leaf area (d).

110

Number of tree and shrub species showing significant variation


in the post-emergent measurements compared to the control for
shoot length (a); radicle length (b); leaf number (c); leaf
area (d).

112

Number of herb species showing significant variation in the


post-emergent measurements compared to the control' for
shoot length (a); radicle length (b); leaf number (c);
leaf area (d).

114

Number of vine species showing significant variation in the


post-emergent measurements compared to the control for
shoot length (a); radicle length (b); leaf number (c);
leaf area (d).

116

13.

14.

15.

16.

17.

18.

19.

20.

21.

22.

xiii

23.

24.

25.

26.

27.

28.

29.

30.

31.

32.

33.

34.

Total number of tree, shrub, vine and herb species showing


significant variation between treatments for each growth
parameter.

116

Number of tree and shrub species showing significant


variation between treatments for each growth parameter.

117

Number of herb species showing significant variation


between treatments for each growth parameter.

118

Number of vine species showing significant variation


between treatments for each growth parameter.

118

Hand TSs of the (a)cineole and (b) camphor chemo-types


of camphor laurel stained with Sudan III showing
possible oil globule variation later identified as not being
related to chemo-type.

140

Cumulative percentage frequency for oil globule length


in the leaves of nineteen street trees of the two chemo-types
of camphor laurel.

142

Cumulative percentage frequency for oil globule length in


the leaves of three wild trees of the two chemo-types of
camphor laurel.

142

Sorted measurements of *C. camphora seedlings below


the chemo-types of *C. camphora and 'control' vegetation
across all sites sampled for: (a) shoot lengths; (b) leaf numbers;
(c) leaf areas.

156

Sorted measurements of *L. lucidum seedlings below the


'camphor' chemo-type of *C. camphora and 'control' vegetation
for all sites sampled for: (a) shoot lengths; (b) leaf numbers;
(c) leaf areas.

157

Sorted measurements of G. semiglauca seedlings below the


'camphor' chemo-type of *C. camphora and 'control' vegetation
across all sites sampled for: (a) shoot lengths; (b) leaf numbers;
(c) leaf areas.

159

Sorted measurements per 150mm of stolon for O. aemulus below


the 'camphor' chemo-type of *C. camphora and 'control' vegetation
for: (a) inter-node lengths; (b) leaf numbers; (c) leaf areas.

161

Sorted measurements of all sampled species other than


*C. camphora found below the 'camphor' chemo-type of
*C. camphora and 'control' vegetation across all sites for:
(a) shoot lengths; (b) leaf numbers; (c) leaf areas.

163

xiv

LIST OF PLATES
Plate No.

1.

Title

Page

Fungal infection of camphor laurel at Uki and Casino, NSW of:


(a) branch-lets; (b) trunk; (c) branches.

177

LIST OF APPENDICES
A1

Germination of the Trees and Shrubs

A2

Germination of the Vines and Climbers

A3

Germination of the Herbs

A4

Vascular Plant Response Groups

A5

Non-linear Regressions -Trees and Shrubs

A6

Non-linear Regressions Herbs

A7

Non-linear Regressions Vines

A8

Modeled data from the Richards Function

A9

Results of the Deviance Tests

A10

Residual Plots of the Deviance Test

A11

T-test Comparisons of the Glasshouse Data

A12

Summary Effect Tables for the Glasshouse Treatments

A13

Non-linear Regressions Algae

A14

Environmental Site Attributes Field Assessment

A15

T-test Comparisons of the Field Data

xv

LIST OF ABREVIATIONS
General
Anon.
ed. (s)
e.g.
et al.
etc.
i.e.
no.
pers. com.
pers. obs.

anonymous
editor(s)
for example
and others
and so on
that is
number
personal communication
personal observation
registered trade mark
copyright

Location and direction


E
EAL
Qld
NSW
S
SCU

east
Environmental Analysis Laboratory
Queensland
New South Wales
south
Southern Cross University

Units of measurement
C
cm
g
h
ha
km
mg/L
m
m2
mm
mm2
MPa
mL
nmol
pH
ppm
s
m
mol

degree Celsius
centimetre
grams
hour
hectare
kilometre
milligrams per litre
metre
square metre
millimeter
square millimeter
megapascal
milliliter
nanomole
a measure of hydrogen ion concentration
parts per million
second
micron
micromole
xvi

Technical
ANOVA
C
CI
C/N
CO2
C-S-R
df
F
FPC
GC(d)
GC-MSFID
GIS
HCl
LS
MAP
n
(NH2)2CO
NH4H2PO4
p
r-K
RIP
REPIL
r2
R
S
S-C
sd
sp.
spp.
TN
TS
TP
UV
%
<
>

analysis of variance
competitor
confidence interval
carbon nitrogen ratio
carbon dioxide
competitive, stress tolerant, ruderal
degrees of freedom
F ratio
foliage projective cover
gas chomatograph(ed)
gas chromatography-mass spectroscopy flame
ionization detector
geographical information system
hydrochloric acid
longitudinal section
mono-ammonium phosphate
size of population
urea
mono-ammonium phosphate
probability
ruderal, stress tolerant
ribosome inactivating protein
repeated mean liklihood
coefficient of determination
ruderal
stress tolerator
stress-tolerant competitor
standard deviation
species (singular)
species (plural)
total nitrogen
transverse section
total phosphorus
ultra-violet
percentage
less than
greater than
less than or equal to
greater than or equal to
introduced plant

xvii

- Chapter 1:Introduction
1.1

Study Background

In many of the regenerating forest remnants and previously cleared land in the northeastern New South Wales which remain outside relatively undisturbed conservation
areas, camphor laurel (*Cinnanomum camphora (L.) Nees and Eberm.) has become
an obvious intrusive exotic species. The trees presence is increasingly presenting
management as well as ecological problems for land owners, plant regenerators and
government authorities. These issues have included: control of the tree in regeneration
works (Dunphy 1991), roadsides (Scanlon 1998, 1999; Stubbs et al. 1999), streamsides, pastures and farms (Stubbs et al. 1999), the economic potential of the tree and
its varied use (Stubbs et al. 1999), possible influence on soil erosion and water quality
(Scott 1999), and whether the tree may be interacting with fauna and flora in an
'adverse' or even 'supportive' manner (Date and Recher 1989, Date et al. 1991, Date
et al. 1992, Schenk and Wallace 1996, Scott 1999, Scanlon et al. 2001, Neilan 2004).

In November 2003, the NSW Scientific Advisory Committee made a preliminary


determination to reject a proposal to list camphor laurel as a Key Threatening Process
under Schedule 3 of the Act, which was provided through Part 2 of that Act (Adam
2003, 2004). The proposal was rejected in early 2004 due, among other things, to
lack of evidence of the alleged toxic effects of the species. The committee found that:

at this time, there is no evidence that camphor laurel (Cinnamomum camphora)


through toxic exudates and leachates in the field situation poses a demonstrable
threat to two or more listed species, populations or endangered ecological
communities, or is likely to cause species, populations or ecological communities to
become threatened. (Adam 2004).

However, the NSW Scientific Advisory Committee supported the declaration of the
tree as a noxious weed in the local government areas of Ballina, Blue Mountains,
Byron, Copmanhurst, Grafton, Hornsby, Ku-ring-gai, Kyogle, Lismore, Maclean,
Pristine Waters, Richmond Valley, Ryde, Tweed and Willoughby, because of its
invasive nature which is presenting major land and resource management issues
(Adam 2004).

To understand the weed status and impact of this tree on Australian ecosystems it is
important to assess what adverse or supportive interactions may be occurring as
this will assist in assessing the current ecological position of camphor laurel and the
possible role of allelopathy in the species overall ecology. A 'supportive' influence
for plant species where habitat might be enhanced has been identified by Schenk and
Wallace (1996) and Neilan (2004), in that high plant diversity was identified in
camphor laurel dominated vegetation of the region. Neilan (2004) suggested that
camphor laurel trees do provide habitat which results from structural change and an
increase in plant diversity of the assemblage through the provision of a suitable
microclimate for the recruitment of many mature phase rainforest plant species and
frugivorous avifauna. Schenk and Wallace (1996) noted that the camphor laurel
assemblages appear to be behaving in a manner similar to a wet sclerophyll forest of
Lophostemon confertus, in that they contain many regenerating native rainforest tree
and shrub species. Date and Recher (1989), Date et al. (1991) and Date et al. (1992)
also commented on the food source provided for frugivorous pigeons in the region, as
a result of the abundant and lengthy fruiting of camphor laurel trees, and the increase
in the numbers of these birds as a result of the spread of the tree.

Coutts-Smith and Downey (2006) identified that alien weed species are the second
greatest threat to biodiversity reduction in New South Wales after habitat loss, placing
camphor laurel as the seventh most significant weed threat to biodiversity after
*Lantana camara, *Chrysanthemoides monilifera, *Rubus fruticosus, *Pennisetum
clandestinum, *Cytisus scoparius and *Ageratina adenophora. With camphor laurel
these authors identified that 6 plant species, 1 animal species and 4 ecological
communities are currently threatened due to the trees invasion. The threatened plants
include Angiopteris evecta, Davidsonia jerseyana, Davidsonia johnsonii, Elaeocarpus
williamsianus, Endiandra floydii, and Endiandra muelleri subsp. bracteata; while the
2

ecological communities threatened are the Illawarra subtropical rainforest, subtropical


coastal floodplain forest, swamp sclerophyll forest on the coastal plain and swampoak floodplain forest. The only animal identified as being threatened by the tree is the
Coxens fig-parrot (Cyclopsitta diophthalma coxeni). Swarbrick (1991) argued that
invasive trees are the most damaging to natural ecosystems due to their ability to
dominate the structure of the vegetation, referring to these plants as canopy-dominant
weeds. These can become monospecies dominant, and may replace or overgrow the
existing canopy, thereby altering the nature and functioning of that ecosystem to a
significant extent (Swarbrick 1991). It is important to note that aggressiveness, lifeform, the ability of the weed to act as point source for spread, method of dispersal,
and reproductive ability are important characteristics to consider when assessing the
invasive potential of a plant (Swarbrick 1991). This is reminiscent of how Grime
(1979) viewed plant strategies, further discussed in this chapter. Firth (1979) and
Dunphy (1991) identified the invasiveness of camphor laurel in north-eastern NSW
and its serious nature as an environmental weed in the region, particularly on the
edges of remnants, below individual trees, in regrowth, and on degraded farmland.
What also emphasizes this fact is that seed originating from the soil seedbank and
neighbouring trees results in an inevitable reinfestation following removal (Dunphy
1991). Batianoff and Butler (2002) found that camphor laurel is in the top ten most
invasive species in south-eastern Queensland, scoring 4.8 on panel data of
invasiveness with 5 having the highest invasive nature to 3 being moderately invasive.
Of the 200 invasive plants indicated to be present in that region, the only other trees
considered to have a higher or equal ranking than *C. camphora was *Celtis sinensis
at 4.9 and *Schinus terebinthifolius at 4.8 on that scale (Batianoff and Butler 2002).
The behaviour of this tree in north-eastern NSW is also very unlike that in its natural
habitat. In mainland China, Chen et al. (2004) indicates that it is a rare tree, occurring
often as individuals in mesic subtropical forests. Miyawaki (1983) in Japan, found
that it is a mid-successional species which grows well in light-open pine woods and is
rarely found as a tall tree in broad-leaved evergreen forest. The presence of this tree in
pine woods in Japan does suggest that the species can tolerate fire prone environments
to some extent, such as exist in Australia. The question posed by this is: Why is the
trees behaviour so different in north-eastern NSW when compared to Asia?

It has been suggested that an 'adverse' influence on plant growth or succession may
result from this tree through competitive advantage throughout its latter growth stages
(Firth 1979, Stewart 2000, Scanlon et al. 2001), large seed numbers (Firth 1979,
Stewart 2000, Scanlon et al. 2001), few natural predators of significance (Firth 1979,
Stewart 2000, Scanlon et al. 2001), rapid spread (Firth 1979, Scanlon et al. 2001), the
formation of mono-cultures (Firth 1979), biomass dominance (Firth 1979, Neilan
2004), association with other weeds (Firth 1979, Schenk and Wallace 1996, Scanlon
et al. 2001, Neilan 2004), destabilization of stream-banks and steep slopes (Scanlon et
al. 2001), the trees longevity (Firth 1979, Valder 1999, Pakenham 2003) and the
blockage of succession (Firth 1979, Scanlon et al. 2001). Allelopathy has also been
indicated by some authors as being a possibility for this tree (Firth 1979, Chou et al.
1989), particularly because volatile terpenes have been identified as being present
(Stubbs and Brushett 2001, Stubbs et al. 2004), in addition to phytotoxic phenolics
(Chou et al. 1989).

According to Cronk and Fuller (1995) the threats to natural ecosystems as a result of a
plant invasion may be varied. These include: replacement of diverse systems with
single species stands, invasion leading to plant extinction or a threat to the native
fauna, and alteration of soil chemistry, geomorphological process, hydrology or the
fire regime. Plant invasion is indicated to progress through various stages which
include an introduction, naturalization through the formation of self-sustaining
populations, facilitation by dispersal agents, spread as a result of successful
recruitment in new locations, interaction with animals and plants, and finally
stabilization which may or may not involve dominance of the vegetation (Cronk and
Fuller 1995). These are important processes to consider in assessing the current
ecological position of this plant in relation to its invasion cycle in the region. It
provides supportive information on the plants adverse or supportive role e.g. a
plant in the early stages of invasion may appear to have little influence, while the
same plant in its latter invasion stage may have obvious interaction with organisms.

Dispersal and establishment of camphor laurel in the north-eastern region of NSW


have been rapid since the first recorded introduction in the region in 1854 (Stubbs et
al. 1999). This is attributed to efficient dispersal by many bird species, and favourable
climatic and ecological conditions for establishment, recruitment and spread (Firth
4

1979). Recruitment appears to be most successful in regenerating forest or open


habitat where browsing and seed predation are reduced in frequency, and moisture
and nutrients remain relatively high, particularly on the Ferrosol (Krasnozem) soils of
the region (Firth 1979, Stewart 1995). Camphor laurel often dominates either as an
extensive intrusion into native vegetation or as monotypic-like stands. The biomass
according to Neilan (2004) exceeds most other exotic weeds and regenerating native
trees by as much as four times the combined basal area of both the native and other
exotic trees. In other words, this is a canopy-dominant weed as referred to by
Swarbrick (1991). The tree is not only confined to the valley locations where Ferrosol
soil is frequently found but it is also observed on various soil types, and on ridge-tops
and drier habitats of the upper slopes (Firth 1979, Stubbs et al. 1999), such as
regenerating dry rainforest and dry sclerophyll forests of the Lismore area (author,
pers. obs.). In these marginal habitats it is observed that tree size is often reduced and
the invasion cycle appears slower over time (Firth 1979). However, highly intrusive
stands of the tree are seen to be forming in some of these drier, more marginal habitats
in the hinterland and individual trees of reduced vigour can also be observed on the
plinths formed at the base of Melaleuca quinquenervia in coastal wetlands, and in
open littoral rainforest habitats (author, pers. obs.). Many regenerating forest areas
throughout the Tweed and Richmond regions, which are not as yet dominated by a
camphor laurel canopy, are seen to be experiencing various stages of dispersal into,
and recruitment by, camphor laurel (Schenk and Wallace 1996). This is evident from
the common presence of germinating camphor laurel seedlings, recruiting saplings
and trees of various girths in a wide variety of open hinterland habitats such as along
fence-lines (Firth 1979), below exposed trees and in regenerating forests (Firth 1979,
Stewart 1995, Schenk and Wallace 1996) and open pastures (Firth 1979), particularly
where grazing is reduced (Firth 1979). Neilan (2004), found very few seedlings on the
forest floor in the twenty-four camphor forests studied, but this may have been due to
the presence of browsing cattle noted in many of the study locations in addition to
decreased light, moisture and nutrient levels under camphor laurel trees. Existing GIS
mapping by various local Councils and State government authorities1 has identified
mature or near mature stands of the tree. However, not many of the areas currently
1

Mapping of Camphor forests in the region has been undertaken by many local councils in northern
NSW (Scanlon et.al 2001), NSW National Parks and Wildlife Service, and the Department of
Infrastructure Planning and Natural Resources.

mapped have included the early stages of recruitment due to mapping being largely
based on aerial photography and/or satellite imagery which identify the upper canopy
characteristics with limited ground identification of under-storey attributes. In view of
the observation of large numbers of camphor laurel seedlings below the canopy in
various regenerating vegetation assemblages of the region, plus the inability of
mapping to provide a more extensive review of the regeneration locations, the
presence of the species is much more extensive than mapping identifies.

The trees invasion has resulted in extensive expansion into various forest types in the
region and seedlings and saplings are observed to be recruiting across open and less
often closed vegetation formations (author, per. obs.). This suggests that camphor
laurel has reached the second last and third last stage of the invasion i.e. spread and
interaction with animals and plants, as it has successfully become naturalized as self
sustaining populations in many locations. The invasion has not only occurred on the
nutrient rich and moister Ferrosols but also in many other, often drier marginal
habitats (Firth 1979). It identifies that the species is in contact and therefore
interacting with a broad range of ecotypes and organisms. It is this stage in the trees
invasion cycle which requires assessment for any possible threat to the plant
communities invaded. The tree does not at first visually appear to be a fitting in
invasive where the presence of the species does not produce high competition or
dominance, but may be more conforming to a pushing out strategy where it actually
competes aggressively and dominates extensively resulting in a lesser ability of the
native vegetation to grow or regenerate (Cronk and Fuller 1995). Is this visual
observation really an accurate assessment of how camphor laurel is interacting with
the flora of the region i.e. does it fit-in or actually push-out as the passing visual
observation may indicate?

Vegetation survey work by Schenk and Wallace (1996) and Neilan (2004) may
indicate that some 'fitting-in' of the species may be occurring as the camphor laurel
stands do provide habitat conducive to the recruitment of mature-phase rainforest
plants (Neilan 2004), as well as many species associated with wet sclerophyll
communities (Schenk and Wallace 1996). In the vegetation survey by Schenk and
Wallace (1996), 179 regenerating forest remnants were surveyed across the Tweed
Valley using rapid assessment methodology. Data gathered on diversity and
6

abundance of plant species in camphor laurel and native dominated regeneration


identified a relatively high diversity of native tree and shrub species in the camphor
laurel forests, and the presence of many rare and threatened species, i.e. 94% of the
thirty-two camphor laurel-dominated plant communities contained rare and threatened
plants, indicating some fitting-in for plant species which are less ecologically robust.
The diversity of tree and shrub species in the camphor laurel forests of the hills
included 27 canopy species and 63 mid-layer species, while the riparian camphor
laurel forests contained 18 canopy species and 46 mid-layer species. However,
camphor laurel stands, although having high plant diversity in some sites, also
displayed a distinct paucity of species at other locations, with some camphor laureldominated sites having few other species including exotic weeds (author, pers. obs).
This has previously been noted by Woodford (1993) who found a difference in the
diversity of species regenerating under a camphor laurel canopy at Victoria Park
Nature Reserve, near Alstonville. One site under camphor laurel contained 10-15
regenerating native species while another site had 65-70 native species. This was
attributed to possible differences in predation of seed and browsing of seedlings at the
two sites, as the light level was indicated to be similar.

In Neilans (2004) more intensive survey of 24 camphor laurel-dominated remnants,


over 200 species of rainforest plants were identified as co-existing in these forests.
These included trees, shrubs, vines, herbs and epiphytes. The study highlighted that
bird dispersal and distance from major seed source areas were important interlinked
factors in the presence and abundance of many plant species in the camphor laurel
assemblages. The study found 113 rainforest plant species likely to be bird-dispersed
in an area of 1.2 ha, and of the 76 juvenile rainforest species recorded, 60 species
were most likely to be bird dispersed. The patterns of recruitment were influenced by
the distribution and abundance of frugivorous birds. Regenerating camphor laurel was
indicated to have a structural complexity which improved upon pasture as habitat for
rainforest organisms and that this complexity provided a microclimate conducive to
the recruitment of rainforest plants. Some elements of structural complexity were
apparent but this was not as well developed as in intact rainforest. There was a low
occurrence of special life forms which are present in rainforest, such as arboreal
epiphytes, trunk climbers, tree ferns and strangler figs. Although thin-stemmed vines
were present, there was a rarity or absence of thick-stemmed lianes in the camphor
7

laurel stands the presence of which would be associated with a rainforest structure.
Palms were also in low abundance and appeared more commonly closest to the
Nightcap Range. Woody debris was also found to be less abundant than in rainforest.
However, canopy closure was two-thirds that of intact subtropical rainforest and
important for the provision of a suitable microclimate for rainforest plants and
animals, particularly the germination and recruitment of mature phase rainforest
seedlings, as well as the suppression of many weeds. The weed *Ageratina spp., was
found to be in greater abundance at 3-15km from the Nightcap Range and was
identified as having an ability to suppress the recruitment of other plants. The basal
areas recorded in the camphor laurel stands were also similar to values recorded in
rainforest although dominated by the presence of camphor laurel. Neilan (2004)
indicated that this structural complexity recorded in the camphor laurel regeneration
sites may develop further with time and would enable the return of many rainforest
species to the Big Scrub region i.e. the area in north-eastern NSW surrounding
Lismore previously occupied by rainforest.

Both the work by Schenk and Wallace (1996) and that of Neilan (2004) identified that
exotic plants contributed little to the overall plant diversity in the camphor laureldominated vegetation but were far more abundant than the native species. Schenk and
Wallace (1996) also found that the camphor laurel tree, in addition to the scandant
exotic shrub, *Lantana camara, and the privets, *Ligustrum lucidum and *L.
chinensis, were intruding into a high number of regenerating vegetation remnants i.e.
26% of the 179 remnants contained weed coverage of 50-100%; 19% with weed
coverage of 20-50%; 32% with weed coverage of 5-20%; and 22% with coverage of
0-5%. Camphor laurel is indicated to be the major component of this weed biomass
when it is considered that 44% of all remnants surveyed had weed intrusion into the
upper canopy of the native trees.

Due to the extensive nature of the mature stands of camphor laurel in north-eastern
New South Wales (Scanlon et al. 2001), the mostly unknown nature of the extent and
location of recruitment, and the trees increasing biomass dominance, these factors
strongly suggest that further management and ecological implications will emerge in
the future. This may become more apparent as the tree expands its presence and range
throughout the region.
8

Neilan (2004) suggests that further research is required on the possible influence of
camphor laurel on plant succession. It is indicated that managing the succession in
camphor laurel-dominated regeneration may be a cost-effective approach to managing
the restoration of rainforest in the Big Scrub region. Successional processors need to
be identified as do the habitat requirements of avian frugivores involved in seed
dispersal in the camphor laurel dominated vegetation (Neilan 2004). These include the
patterns of rainforest regeneration, the factors affecting plant recruitment such as
predation and herbivory, seed rain, and the use of regrowth as habitat by animals
(Bower 2004, Kanowski et al. 2004). Scanlon et al. (2001) also indicate that there is a
limited knowledge of the most likely course of succession in the camphor laurel
regeneration as well as the consequences of management intervention. Early work by
Firth (1979) calls for research into possible biological control of the tree. However,
before this can be implemented the status of the tree in relation to its threat status as
an environmental weed needs to be further understood in addition to a greater
understanding of the general ecology of the tree in context of the Australian
environment.

1.2

Capability Attributes of Camphor Laurel


1.2.1

Brief Physical Description and Recent Classifications

Camphor laurel is a large robust evergreen tree which is capable of exceeding a height
of 40m and a girth of 22m in southern Japan (Pakenham 2003). These are old growth
trees exhibiting signs of upper limb die-back, limb hollows and basal decay. The trees
in Australia are often less than 100 years old, and although not of this size, tend to
over-canopy the slower growing native species early in its life-history, forming dense
dominating stands.

Camphor laurel seedlings grow rapidly into spindly saplings which display at an early
age the efficient light capturing architecture of the vase shape with upwardly angled
branches containing rounded densely packed whorled leaf clusters arranged in a
pyramid like sequence. This shape is maintained in the mature tree with the
development of robust branches, trunk and main roots capable of substantial
epicormic resprouting which can further add to the overall vase shape. The

development of a lignotuber-like structure early in the growth of seedlings in addition


to a large basal swelling of the trunk in mature trees is present possibly for moisture,
nutrient and allelochemical storage. Bark is thick and fissured and has been observed
by the author to heal quickly and with limited decay of the exposed wood. There is
also limited limb drop which contributes to biomass retention. The root system also
possesses massive radiating main roots and dense net-like feeder roots which exploits
nutrients and water in the upper soil horizon. Only a few main roots penetrate deeply
into the soil (author pers obs.).

Camphor laurel belongs in the family Lauraceae. Recent taxonomic descriptions of


the camphor laurel tree appear in Firth (1979), Harden (1992), Liao (1996) and Le
Cussan et al. (2007).

In Australia, Harden (1992) and Le Cussan et al. (2007) recognize the species as
Cinnamomum camphora (L.) J.Presl., a name recognized since 1825, while
Cinnamomum camphora (L.) T.Nees & C.Eberm. is indicated by Chapman (1991) to
be in use since 1831.

According to Liao (1996), two varieties are now recognized in Taiwan. This
taxonomic classification has mostly absorbed the more complex species, variety and
subvarieties listed by Firth (1979) in Taiwan. Previous classifications of: Laurus
camphora L., Cinnamomum camphora (L.) Nees & Eberm. var. nominale Hayata
subvar. hosyo Hatusima. and Cinnamomum camphora (L.) Nees & Eberm. var. hosyo
(Hatusima) Liao are now recognized as Cinnamomum camphora (L.) Presl, Prinoz
var. camphora. Cinnamomum camphora (L.) Nees & Eberm. var. glaucescens
(Braun) Meissn., Cinnamomum camphoroides Hayata., Cinnamomum nominale
(Hayata ex Matsum & Hayata) Hayata and Cinnamomum camphora (L.) Nees &
Eberm. var. lanata Nakai are referred to by Liao (1996) as Cinnamomum camphora
(L.) Presl, Prinoz var. nominale Hayata ex Matsum. & Hayata.

The two varieties are differentiated by Liao (1996) according to the presence of bark
knobs on the trunk in the var. nominale and an absence of knobs in the var. camphora.
Examination of the Australian trees identifies the absence of knobs, indicating that the
variety is *Cinnamomum camphora (L.) Presl, Prinoz var. camphora. according to
10

Liaos classification. However, the classification *Cinnamomum camphora (L.) Nees


and Eberm is used by recent authors in Australia such as Stubbs and Brushett (2001)
and Stubbs et al. (2004).

Various chemo-types of the tree have also been identified at varying geographical
locations in its natural range, identifying genetic variation below the species level.
These are based on the dominating oil present in the leaf and branches and include the
chemo-types camphor, linalool, 1,8-cineole, nerolidol and borneol (7-10) (Shi
et al. 1989, Zhu, et al. 1993,

Zhu et al. 1994, and Lawrence 1995). Initial

identification of chemo-types in Australia by Hirota and Hiroi (1967) and the gas
chromatograph work by Stubbs and Brushett (2001), Stubbs et al. (2004) in addition
this current work, identified that only two chemo-types are so far known to be present
in the region i.e. the camphor chemo-type and the cineole chemo-type.

1.2.2

Geographical Range and Climate

Camphor laurel is naturally distributed throughout the subtropical regions of southern


China (Grieve 1931, Liao 1996, Chen et al. 2004) in the provinces of Jiangxi,
Guandong, Guangxi, Hunan, Hubei, Yunnan, Zhejiang, and Fujian (Chen et al. 2004)
nearly to the mouth of the Yang-tse-kiang River in the south (Dewey 1897); as well as
Taiwan, Korea, Vietnam (Liao 1996, Chen et al. 2004), Japan (Grieve 1931, Liao
1996, Chen et al. 2004), Hainan (Dewey 1897) and the Ryukyus islands (Liao 1996).

The natural habitats in which it is found include subtropical forests in southern China
(Chen et al. 2004) and in Japan, light-open pine woods of Pinus merkusii and rarely as
a tall tree in broad-leaved evergreen forest (Miyawaki 1983). In Taiwan var.
camphora is found at low elevation in the north and south while the var. nominale is
endemic to Taiwan occurring in scattered locations in the east and south of the island.

It has been introduced to many other countries including Australia (Firth 1979, Chen
et al. 2004), Florida in the USA (Grieve 1931, Chen et al. 2004), California in the
USA (Grieve 1931), Argentina (Grieve 1931, Huergo and Retamar 1978, Frizzo et al.
2000), India (Grieve 1931, Baruah et al. 1975, Ghosh 1977, Bahaguna et al. 1987,
Balasubramanian 1993), Bangladesh (Sattar et al. 1991), Ivory Coast (Pelissier et al.
11

1995), Cuba (Pino and Fuentes 1998), South Africa (Immelman et al. 1973, Cronk
and Fuller 1995) in the mesic subtropical forests (Immelman et al. 1973), the shores
of the Black Sea (Mikhalevskaya et al.1993), Madagascar (Grieve 1931, De Medicei
et al. 1992), Malaysia (Jantan and Goh 1992), and Sri Lanka, Egypt, southern Europe,
and the Canary Islands (Grieve 1931). It has become an environmental weed in
Australia (Firth 1979, Batianoff and Butler 2002), South Africa (Cronk and Fuller
1995) and Florida (Laessle 1958, Daubenmire 1990).

The current locations where camphor laurel is now growly throughout the World
identifies that the naturalized distribution of this tree is subtropical, with some overlap
into tropical and temperate regions where temperatures are not too high or low for the
species. In its natural range in mainland China the climate is subtropical, humid and
monsoonal, receiving regular typhoons and often having a well-defined dry period in
spring from March to May. In Hebei province, however, which is at the extreme
northern and western limit of camphor laurel distribution in mainland China, there is a
temperate continental climate with dry winters (Chinese Business World 2004). Chen
et al. (2004) found that the long dry periods in China coincide with flowering and
fruiting, reducing fruit numbers due to the dry conditions. He suggested that in
Australia the tree flowers and fruits during a period of high rainfall allowing for high
rates of pollination and fruiting as noted by Firth (1979).

In Taiwan the tree is confined to the lowlands where it reaches 1,200m in the north
and 1,800m in the south (Liao 1996) identifying its response to altitudinal and
latitudinal temperature. Table 1 identifies the climatic features of the various regions
of China where camphor laurel has been found by Chen et al. (2004) to grow. Taiwan
is identified to be the warmest and wettest region and it would be expected that
camphor laurel would appear at lower elevations in the mainland provinces, as
temperature decreases towards the north and west, particularly in view of the
altitudinal variation between the north and south of Taiwan (Liao 1996).

In India, it is cultivated extensively in gardens in Dehra Dun, Saharanpur, Calcutta,


Negiris and Mysore and is capable of growing up to an altitude of approximately
1,300m in the north-west Himalayas (Chopra et al. 1956).

12

Four widespread locations in Australia where camphor laurel is known to be present


as self-sustaining stands are climatically compared in Table 2. These include around
Nowra, where it is identified to be at the southern range of its spread in NSW as a
weed, and the Tweed where it is at its northern range in NSW (Firth 1979); Gladstone
at the northern limit of the weed study by Batianoff and Butler (2002) on the midcoast of Queensland; and Cooktown in far-north Queensland which is the northern
limit of its spread on the east coast of Australia (Firth 1979). Street plantings have
also occurred in Toowoomba and Armidale being cooler high altitude locations in
Australia. However, these trees at higher altitudes have not become invasive due to
frost sensitivity of the seedlings (Firth 1979).
Comparison of the climatic data presented in Tables 1 and 2 for the areas where
camphor laurel is known to grow in China according to Chen et al. (2004), and in
Australia by Firth (1979) and Batianoff and Butler (2002) reveal the following:

annual temperatures vary from -14 to 31 oC in China and Taiwan, while in


Australia they vary from 8 to 32 oC;

the summer growth periods are reversed, as would be expected due to the

shifts in northern and southern hemisphere climate throughout the year, providing
maximum temperatures for growth in China and Taiwan during July, and in Australia
during January;

annual precipitation range from 400 to 5000mm in China and Taiwan where
camphor laurel is distributed, while in Australia it varies from 974 to 1665mm; and

the wettest period in China and Taiwan can extend from winter to early

summer, with a pronounced dry period for several months during spring and summer
in many provinces. This would curtail plant growth in these regions. In Australia,
most precipitation also occurs in summer i.e. around December to January, but the
difference lies is that rainfall continues throughout the summer in Australia, with only
a brief drier period in spring in comparison with China. However, at the extreme
southern end of camphor laurel distribution (as a weed) in Australia, at Nowra, this
drier period extends throughout the spring.

13

Table 1. Temperature and precipitation for the provinces of southern China in which
camphor laurel is naturally distributed, according to Chen et al. (2004).

Province

January
Temperature
o
C

July
Temperature
o
C

Annual
Precipitation
mm

Maximum
Precipitation
Period

Minimum
Precipitation
Period

Hebei

-14 to -2

20-27

400-800

July

Zhejiang

2-8

27-30

850-1700

Jiangxi

3-9

27-31

1200-1900

Hunan

4-8

26-30

1250-1750

June early
July
half in AprilJune
March -July

DecemberFebruary
July-August

Fujian

5-13

25-30

800-900

May-June

Guangxi

6-16

25-29

1200-1800

Yunnan

8-17

11-29

600-2300

AprilSeptember
May-October

Guandong

8-17

27-29

1500

SeptemberNovember
JuneNovember
March-May
OctoberMarch
NovemberApril
June-August

AprilSeptember
Taiwan
13-20
24-29
2000-5000
Throughout
None
year
___________________________________________________________________________________
Note: Provinces are listed from lowest to highest minimum temperature coinciding with north to south geographical location.

source: Chinese Business World (2004)

Table 2. Temperature and precipitation for four Australian locations where weed invasion
resulting from camphor laurel has been recorded by Firth (1979) and Batianoff and Butler
(2002).

Location

January
Temperature
o
C

July
Temperature
o
C

Annual
Precipitation
mm

Maximum
Precipitation
Period

Nowra

16-28

8-17

974

January-July

Minimum
Precipitation
Period

AugustNovember
Murwillumbah
20-30
9-21
1585
DecemberAugustMay
September
Gladstone
22-30
11-22
1021
DecemberAugustMarch
September
Cooktown
24-32
18-26
1665
Dec-April
SeptemberOctober
_____________________________________________________________________________
Note: Locations are listed from lowest to highest minimum temperature coinciding with south to north geographical location.

source: Australian Government Bureau of Meteorology (2004a, b)

This climatic comparison suggests that camphor laurel has the potential to spread to
regions of lower temperature, and lower and higher rainfall in Australia than it now
occupies, but not within the hotter tropical areas of the country or regions receiving
high frost. This indicates that it may be at the fourth stage of a six stage invasion
14

process as defined by Cronk and Fuller (1995) i.e. spread of the species across a broad
region.
1.2.3

The Ecological Strategies of Camphor Laurel

Camphor laurel does exhibit a number of possible adaptations and strategies which
may enable the tree to be an efficient and widespread intrusive or dominant species in
native bush-land of the region.

Following a preliminary assessment of the trees ability to have such an ecological


advantage over other species in north-eastern New South Wales (Schenk 1998),
thoughts on these hypothesized adaptations and strategies have been further
developed and reviewed, providing an ecological profile which addresses the plants
adaptive strategies.

Primary Strategy
Grime (1979) identified that plants respond to competition, stress and disturbance in
various ways, defining these as: competitionthe ability of neighbouring plants to use
the same light, nutrients, moisture and space; stressthe external constraints that may
limit the accumulation of dry matter produced in part or all of the plant; and
disturbancethe mechanisms that limit biomass through the partial or total destruction
of the plant. Using the principles of competition, stress and disturbance, Grime (1979)
was able distinguish three distinct strategies in plants: Primary strategies in the
established phase; Secondary strategies in the established phase and; Regenerative
strategies.

The primary strategy involves competitive ability and stress tolerance enabling
advantages in particular ecological situations (Grime 1979). These involve three
groups of plants:

competitors (C), allowing exploitation of low stress and low disturbance,

stress-tolerators (S), allowing exploitation of high stress and low disturbance,


and

15

ruderals (R), allowing exploitation under conditions of low stress and high
disturbance.

However, several intermediate strategies also exist, referred to by Grime (1979) as


secondary strategies. These strategies also occur in the established phase of the plants
growth cycle and may result from a combination of the primary strategies identified
by Grime (1979). These include competitive-ruderals (C-R), stress-tolerant ruderals
(S-R), stress-tolerant competitors (S-C), and competitive-stress tolerant ruderals (CS-R). Regeneration strategies are dealt with separately by Grime (1979) as it is
indicated that these are distinct from the strategies employed by the mature plant.
Therefore the profile that follows subscribes to this general approach providing a
structured level of comparison, enabling the identification of the possible reasons for
the trees success.

Grime (1979) indicated that competitive ability is a result of a combination of plant


characteristics, being a function of area, activity and distribution of the plant surfaces
through space and time. Specifically these plant characteristics may include the
presence or absence of storage organs, height of the plant, lateral spread, phenology
(density of foliage and its relationship to sugar production), growth rates, response to
stress and damage, and the role played by intraspecific variation in the plants
competitive ability. For camphor laurel these have been identified in brief as:

growth and productivity (rapid growth including high productivity and


biomass accumulation),

storage organs (presence of lignotuber like storage organ, large trunk and
primary roots enabling rapid recovery following stress or disturbance events
and as a possible seedling reserve),

longevity (long lived nature of the tree allowing persistent and long term
dominance),

adaptive ability (genotypic and phenotypic plasticity allowing morphogenetic


adjustment through natural selection from a large gene pool),

16

inhibition (possible allelopathic compounds, moisture and nutrient reduction


below the tree due to shallow, invasive and massive root system, and limited
light levels below the mature canopy), and

response to stress (physiological and morphological adaptations and responses


to low moisture and UV radiation, rapid frost response, fire retardant ability of
the bark and an extensive coppicing ability of the trunk, branches and main
roots).

Productivity
Factors involved in high productivity and biomass accumulation by the camphor
laurel tree are a specialized leaf abscission strategy enabling sugar production
throughout the year with the sugars peaks during periods of seed production and
growth, and the dense compacted nature of the leaf arrangement and architecture
whose shape maximizes photosynthetic area. This provides a growth advantage in the
later stages of growth prior to maturity and at maturity through its biomass dominance
of the site (Firth 1979). Early growth was identified by Firth (1979) as being less
competitive. The observed rapid growth of monotypic-like stands in native
regeneration assemblages of a fairly even age class is visually evident in many
locations. In mature stands, biomass is observed to exceed that of many native and
other exotic tree species on site as indicated by Neilan (2004).

The strategies involved in high productivity of this tree include vernal abscission
enabling higher photosynthetic ability and a highly efficient light intercepting
architecture and leaf arrangement.

As described by Addicott (1978), vernal abscission is a process where leaf fall, new
leaf formation and inflorescence development coincide. This is observed in the bud
formation behaviour of camphor laurel, occurring twice per year. Abscission of the
previous year's leaves occurs with the commencement of the most favourable growing
period when new growth is beginning. It is triggered by development of these new
sinks in the tree i.e. the new leaf flushes, by becoming richer sources of auxin and
other hormones and the older leavers weaken as hormone sources, resulting in
abscission (Addicott 1978). Firth (1979) identified that leaf abscission commences

17

following the advanced formation of the leaf flushes and observed in this research to
occur at the same time as flower formation which is twice yearly in the warmer lower
altitudes of the Big Scrub region. This process of vernal abscission would enable
maximization of sugar production during favourable growth periods as old inefficient
leaves are replaced immediately by new more efficient leaves which are more
photosynthetically active at producing sugars. The tree is, therefore, not only able to
maximize sugar production at fruiting due to the coincidence of new leaf and flower
development, but also able to produce sugars at less efficiency throughout periods
when leaf formation and fruit development are inactive. Firth (1979) observed that
growth does occur throughout the year but is reduced during periods when leaf
formation is inactive. Immediate loss of old leaves at the time of new leaf formation
would provide an advantage in that no period during the year would be without
photosynthesis, with older less efficient leaves replaced regularly during periods of
peak sugar demand.

Temperature Triggered Growth


Yurugi and Nagata (1981) in Japan found that camphor laurel produced growth during
periods of both long day and short day lengths, indicating that bud dormancy is not
induced or released as a result of photoperiod. However, the research clearly
demonstrated that flushing is triggered by temperatures above 18C, establishing that
temperature and not photoperiod is responsible for triggering flushing behaviour. In
Japan it was found that the tree is induced to bud dormancy with lowering
temperatures in autumn, and released from dormancy with winter chilling in that
region (Yurugi and Nagata 1981). This indicates that camphor laurel which grows in
regions where temperature is sustained above 18C will maintain constant growth,
without constraint.
Further work on the influence of higher temperature on the growth of camphor laurel
in Taiwan by Sheu and Chang (2001) indicated that as temperature increases from
between 23-28C to between 28-33C, the growth of seedlings measured as net height
and net weight also increases. However, when C02 concentration increases from
350ppm to 700ppm productivity decreases at the higher temperatures. This indicates
that camphor laurel productivity is limited to some extent by temperature and
excessive concentrations of C02.

18

In a comparative study of the influence of low temperatures on the development and


growth of buds in both *C. camphora and *C. japonicum in Russia, by
Mikhalevskaya et al. (1993), the former species was found to have a shorter period of
apical meristem dormancy, and an earlier initiation and later cessation of shoot
growth. Frost resistance was also found to be positively correlated with the length of
bud dormancy between these two species, with *C. camphora the more frost
susceptible as a result of these earlier and later periods of growth. Mature *C.
camphora of an age of 74-76 years was susceptible to frost of -8C with 70% damage
to leaves, and -10C with 100% damage to leaves and *C. japonicum at -14C had 1020% damage, and at -16C, 100% damage.

Canopy Architecture
Tree architecture too may be important in the maximization of photosynthetic area
available for light interception. The canopy of camphor laurel is dense with leaves
closely arranged in pseudo-whorls or alternately on the branchlets which are clustered
tightly in the canopy, allowing little entry of sunlight below, particularly in a mature
stand. Additionally, the response of the tree to removal of the apical meristem which
might occur as the result of fire or browsing is the initiation of bud growth close to
ground level in many cases, producing radiating trunks which develop a vase-shaped
architecture, eventually forming the distinctive rounded and densely interlocked
canopy of the mature tree. In either case, it appears that the use of available
photosynthetic area is maximized (author, pers. obs.).

Limited light levels below the mature canopy have been noted by Firth (1979) and
Stewart (1995). Canopy closure may also be influencing some light-demanding
regeneration species such as Eucalyptus spp., Acacia spp., Allocasuarina spp.,
Baeckia spp., Leptospermum spp. and Callistemon spp. amongst others. Schenk and
Wallace (1996) and Neilan (2004) found that rainforest plants do germinate and grow
beneath the camphor laurel canopy, particularly mature phase bird-dispersed species
(Neilan 2004). Light levels therefore could be seen as a significant factor in
influencing succession in the camphor laurel regeneration.

19

Root Architecture
Observation of soil dryness is evident below the mature camphor laurel canopy (Firth
1979). Examination of the root architecture in roadside cuttings reveals large and
extensive radiating roots close to the soil surface (Firth 1979). Hadington and Johnson
(1979) also examined this tree for its root invasiveness, identifying that it was one of
the six most aggressive root invaders in developed areas in Australia, invading sewer
pipes and also being able to lift paths and damage wall foundations. Clearly this
identifies the ability of the roots to aggressively exploit moisture reserves in the soil
and contribute to soil dryness.

The observed dryness and possible resulting reduction in nutrient availability may
discourage, through higher below ground root competition, the germination of plant
species, particularly some mesic rainforest plants that rely on moist conditions for
germination and have limited seed longevity once dispersed (author, per. obs.).

Inhibition
Inhibition in camphor laurel is identified to result from the possible presence of
allelopathic compounds, moisture and nutrient reduction in the upper soil horizon as a
result of a massive shallow root system and low light levels below the mature canopy.

A complex allelopathy may be present which may inhibit or prevent the germination
or growth of other plant species. Rice (1984) defined allelopathy as being distinct
from plant competition in that it involves biochemical interactions, but importantly it
can involve both inhibitory and stimulatory responses from the same compound.
Inhibitory allelopathy has been indicated by Rice (1964, 1965 and 1984), Muller et al.
(1964), Whittaker (1970), Muller (1974) and Leicach et al. (2009) for the essential
oils cineole and camphor, which have been chromatographically assessed by Stubbs
and Brushett (2001) and Stubbs et al. (2004) as being in relatively high concentrations
in the camphor laurel tree, in addition to twelve other volatile oils. Muller et al.
(1964), Muller (1974) and Rice (1984) suggest that these volatile compounds may
either be lost to the atmosphere or even adsorbed on to soil particles. However, Firth
(1979) recognized that soil release of aromatic compounds was occurring, as a
distinctive odour was found to be present in soil samples taken from the vicinity of
camphor laurel trees, although no effect on the growth of plants in this soil resulted in
20

a glasshouse trial. Rice (1984) discussed the complex nature of allelopathy which not
only involves a complex and varied group of biochemical compounds and pathways,
but is also due to variation in effect due to their concentration, influences due to
varying environmental and soil conditions, and variability in species response. He
suggested that it is possible that any effect observed due to allelopathy would not only
be due to the oils themselves but due to a complex set of these interacting factors. The
oils identified by Stubbs and Brushett (2001) and Stubbs et al. (2004), were used to
establish differences in intraspecific variation and therefore may not be the only
compounds which may have allelopathic activity in this tree.

Apart from the inhibitory influence of the oils cineole and camphor under laboratory
test conditions, the leaf extracts of the camphor laurel tree have been identified as
active on the germination of some seeds by Firth (1979), Johns (1994), and Chou et
al. (1989), although it has not been fully assessed as an 'operating field component' in
the general ecology of the tree. Scott (1999) visually suggests that allelopathy in this
tree may not be effective as seedlings under camphor laurel trees, once the tree is
removed, do grow vigorously; indicating light, moisture and nutrients are responsible
rather than allelopathy. In a study by Paul et al. (2010) on the influence of soil
variation on the seedling growth of rainforest pioneer species in pasture, rainforest
and camphor laurel dominated vegetation, found that growth rates were reduced by
25% in soils below camphor laurel. The study concluded that reforestation pathways
are affected through this influence on seedling growth by altering the physical and
biochemical properties of soils.

An alpha selection resulting in an altered plant succession may possibly occur if direct
allelopathy is identified as active in the field situation for this tree. This would result
in changes to plant succession with those species more tolerant of the allelopathic
effect or stimulated by it becoming more dominant in the succession. However, this
requires further testing to verify whether allelopathy is significant in the ecology of
the species and if any differences in effect may exist between the two chemo-types.
Further detail is provided on allelopathy in camphor laurel later in Section 1.3.3.

It is important to note that allelopathy cannot be viewed in isolation as many


interacting factors such as light, temperature, moisture, soils, predation, herbivory,
21

genetic variation, and interaction with organisms are active, and contributing to the
success of this tree. Allelopathy therefore, must be placed in context with these other
interacting factors, enabling the establishment of the possible role allelopathy has in
the success of the species in north-eastern New South Wales i.e. how important is
allelopathy in the ecology of the tree?
Storage Organs
Storage organs such as a lignotuber-like basal swelling, large trunk diameter to height
ratio and enlarged primary roots are visually evident. This provides for the storage of
moisture, carbohydrates and allelochemicals giving further competitive advantage to
the tree. Even in seedlings, Firth (1979) identified the presence of a root swelling
located at ground level, which was assumed to be a food storage organ. Observations
of mature trees for this study also found that a massive swelling at ground level occurs
in some trees, particularly those that have had the primary trunk removed or damaged
as a result of fire, browsing or disease. New buds arise from this swelling in profusion
enabling rapid re-growth following such events. The swelling appears to resemble the
lignotuber of Eucalyptus spp. which plays a similar role following fire events (author,
pers. obs.). This indicates that the lignotuber-like swelling is not only functioning as a
storage organ but also has a regenerative function through epicormic re-sprouting.

Longevity
Longevity of the tree is also an important factor in this species competitiveness.
Recent publications and websites have indicated that some trees in China are reputed
to be 700 years old (Valder 1999) to 2,000 years old (Peoples Daily Online 2004). In
Japan some trees are said to be 1,000 years old (Japan Broadcasting Corporation
2004), 1,300 years old (Atsutajingu 2004) and 2000 to 3000 years old (Pakenham
2003). In Australia Coutts-Smith and Downey (2006) indicate that the tree can survive
for periods of greater than 400 years. Although these ages may be disputed due to lack
of scientific aging techniques, it does identify that once a camphor laurel tree is
established, it is present for a considerable period of time. This would enable the
continual exertion of the species on a plant community over time, through extended
occupation of the available niche. It would also provide for a continual and
exponential production of seed as recruited generations mature and enlarge their
canopies.

22

Adaptive Ability
Camphor laurel has a high adaptive ability through possessing genetic variability
across its natural range. This variability provides phenotypic and chemotypic
plasticity. Grime (1979) indicates that such variability in a plants plasticity enables
rapid morphogenetic adjustments to various ecological situations. In a forestry
genetics trial of camphor laurel, a high genetic variability across mainland China was
identified on the provenance and family level for seedling height, ground diameter,
branching height (Xiaohua et al. 1999, Xianghua and Liqing 2001) and freeze injury
(Xiaohua et al. 1999). The variance analysis performed by Xiaohua et al. (1999)
demonstrated a highly significant difference among the provenances, but less so at the
family level. It was also found that a highly significant relationship exists between
seedling height, ground diameter and branching height, but no significant relationship
between freeze injury and these traits.

Further work in China by Huadong et al. (2000) found a highly significant difference
in seedling biomass among provenances and a highly significant relationship among
nine traits which included seedling height and ground diameter. Ground diameter of
the seedling was found to be a major factor related to seedling biomass.

Xiahua et al. (2001) also undertook a genetic variation trial on a provenance level
and family level for bud germination and this was found to be highly significant.
Early, mid and late bud germination varies more significantly at the provenance level
than at the family level, and a highly significant relationship exists between these
three bud germination times. Early bud germination and middle bud germination
displayed a highly significant relationship to annual mean temperature, January mean
temperature (the coldest month in China), extremely low temperature, as well as the
number of dry months. Latitudinal variation was also found to have an extreme
negative relationship with early and mid bud germination.

Five chemo-types have been so far identified across its natural range, based on their
volatile oil composition (Shi et al. 1989, Zhu, et al. 1993, Zhu et al. 1994, and
Lawrence 1995) identifying further plasticity in chemical composition i.e. variation
below the species level.

23

This work in Asia demonstrates that camphor laurel possesses not only a high
chemotypic differentiation across its natural range, but also a high genotypic
differentiation found to be expressed in its seedlings from various mainland
provinces.

Heywood (1967) indicated that many organisms of the same genotype can produce
various phenotypes as a result of development in various environments and referred to
this as phenotypic modifications or ecads, which can be expressed as either overall
plasticity or variations in parts or organs of the plant. This means that phenotypic
plasticity is a property specific to individual characters in relation to specific
environmental factors, but can also vary according to the stage of growth. These ecads
can be difficult to establish as differences in effect due to environmental modification
and differences due to genetic variability may appear similar. However, this can be
overcome through artificial modification and control of climate under glasshouse
conditions during a growth trial (Heywood 1967). The work on the camphor laurel
provenances in China did use glasshouse growing conditions and established that the
variation in the features of seedling height, ground diameter, biomass, freeze injury,
and bud germination response is due to genetic plasticity resulting from variability in
the tree from one province to another in China.

However, phenotypic plasticity also occurs in this tree. This is shown in variation in
leaf size on individual trees and has been observed to vary even on individual
branchlets (author, pers. obs).

The chemo-types also appear to be indistinguishable using external morphological


characters. However, physical differences in internal morphology of the chemo-types
may be worthy of further research, as internal morphology of the petiole is distinctive
in camphor laurel compared with other species in the genus Cinnamomum
(Balasubramanian et al. 1993), containing distinctive bands of collenchyma below the
epidermis and an enclosing abaxial band of thick sclerenchyma tissue around the
phloem, with oil glands and mucilaginous cells in the parenchyma ground tissue.

24

Slow Litter Decomposition


Shima and Chiba (2002) undertook a study of the changes in biological nitrogen
fixation and organic decomposition of leaf litter under camphor laurel secondary
forest in Japan for a ten month period. Findings from this study revealed that the leaf
litter of this species decomposes slowly and that: acetylene reduction activity (ARA)
is low, being less than 1 nmolg-1h-1, peaking briefly to 5 nmolg-1h-1; carbon to
nitrogen ratio (C/N) falls rapidly from near 58 to near 22; benzene-ethanol soluble
fraction falls linearly from 15% to 4% of dry matter; water soluble fraction falls
rapidly from 20% to 8% of dry matter; hemi-cellulose increases rapidly from 20% to
28% dry matter then falls rapidly again to 20%; cellulose decreases slowly from 12%
to 9% of dry weight; and lignin content increases linearly from 30% of dry weight to
56%. With these data, Shima and Chiba (2002) demonstrated that nitrogen fixation
by camphor laurel leaf litter is minimal, and that other species are more effective at
fixing nitrogen in this way than this particular tree. Thus nitrogen release from
camphor laurel leaf litter may not be an important factor in any possible nutrient
inhibition. However, the slow decay of litter and the thick accumulation below the
canopy noted by Firth (1979) and Neilan (2004) may discourage some seed
germination particularly of small-fruited species, or give a growth advantage to those
species which are large fruited, shade tolerant and not influenced in an inhibitory way
by possible allelopathic release from the litter.

Stress Tolerance
Camphor laurel responds rapidly to stress and damage through its ability to limit
moisture loss and the influence of UV radiation, the fire retardant ability of its bark,
and the coppicing ability of its trunk.

Adaptations to moisture stress and protection from UV radiation may occur through
possible leaf adaptations such as waxy cuticle, stomata position or function to
reduce moisture loss, enclosed buds, anthocyanin pigment and perhaps specialised
adaptations in water storage within the plant such a large main trunk which may be
multi-stemmed, lignotuber like swellings of the lower trunk, and large primary roots
and branches (author, pers. obs.).

25

Balasubramanian et al. (1993) compared the petiole anatomy of four species of


Cinnamomum i.e. *C. camphora, *C. macrocarpum, *C. tamala and *C. zeylanicum.
They identified that *C. camphora was distinctly different to the other species studied
in that it has a thick hypodermal collenchyma of four to five cells thick and a crescent
shaped vascular strand surrounded by a thick and continuous zone of abaxial
sclerenchyma. These features were absent in the other species studied. However, all
these species were found to possess a uniseriate and slightly cutinized epidermis. This
may be an indication of the development of some sclerophylly in the leaf of camphor
laurel, although further leaf sectioning is required to verify this.

Research in South Africa by Vander Willigen et al. (2000) on the xylem


characteristics of *C. camphora in comparison to a number of other native tree
species which grow in subtropical mesic forest of the region, identified that this tree
showed the highest vulnerability to cavitation. It lost conductivity over a narrow water
potential range, with 50% loss in conductivity as water potentials approached 1 MPa,
with the water potential gradient gradually decreasing as transpiration increases. This
vulnerability does identify the drought sensitive nature of this tree and suggests a high
intolerance to salt. However, *C. camphora does appear to lose considerable
conductivity during the day without influencing transpiration rates to any great extent,
and that xylem pressure does return to normal overnight indicating overnight refilling.
This was not shown in the other species studied. The conclusion drawn for the study
was that maximum xylem-specific conductivity and vulnerability to cavitation are
genetically determined, and the leaf area supplied by the trees hydraulic pathway is
phenotypically plastic being influenced by the environment.

Response to Physical Damage


Fire may also play a role as a constraint in the forests of the region, particularly on the
drier north facing slopes which are more prone to sunlight exposure. In Florida,
camphor laurel communities were found to be highly invasive when fire is removed
(Laessle 1958, Daubenmire 1990). In the local area, fires may have the effect of
destroying seed, seedlings and small saplings which have not developed a consistently
thick bark layer near ground level (author, pers. obs.). However, Floridata (2004)
indicate that the wood is hard to burn, identifying that mature trees are more resilient

26

to forest fires. The bark of the camphor laurel tree heals quickly with little damage to
the underlying meristematic tissues and little wood decay (author, pers. obs.).

Firth (1979) noted that coppicing occurs along the trunk where grazing or fire
removes the upper limbs in young trees which possess minimal bark coverage and
protection. Physical damage to the branches, trunk or roots stimulates bud growth
arising from previously dormant epicormic buds below the bark surface. This
coppicing strategy may not only be efficient for the interception of available sunlight
but may allow for rapid regeneration following a stress event. The famous camphor
laurel trees of Hiroshima which regrew rapidly following the atomic bomb blast in
1945 (Hiroshima City-Business Placement 2004) demonstrate the recuperative ability
of this tree following severe damage.

Dewey (1897) found that mature trees in Tokyo are subjected to 70-80 nights of frost
and temperature of -11 to -9C and seedlings grown in America can withstand brief
temperatures as low as -12C. Firth (1979) found that trees growing in Armidale and
Toowoomba can withstand temperatures of -9C and occasional snow falls, but have a
low establishment of naturalized trees, which was partly due to frosting. He concluded
that camphor laurel has a limited tolerance to frost in its seedlings, not being able to
survive temperatures below -12C, and found that -5C was enough to cause leaf
browning in seedlings.

Bregvadze et al. (1975) compared the activity of growth regulators in the leaves of
*C. camphora and *C. japonicum when subjected to frost. In *C. camphora there was
a low activity of auxins during winter with no activity in *C. japonicum, while the
activity of inhibitors was lower in *C. camphora, indicating a lower frost tolerance for
camphor laurel.
Mikhalevskaya et al. (1993) in experiments with frost damage on leaf development of
camphor laurel trees along the Black Sea, found that although the apical meristem was
damaged as a result of lengthy periods of growth under cold conditions, the tree
recovers rapidly through the development of lateral sylleptic shoots, but the
comparative species, *C. japonicum was unable to achieve this. This study
demonstrated that *C. camphora produced these buds earlier, at a more rapid rate and

27

in higher number following frost damage events at -10 and -14C, indicating that
mature camphor laurel although significantly damaged by frost at these temperatures
can recover rapidly.

Adaptations for stress tolerance and a rapid response to disturbance are further lifeattributes which provide competitive advantage for camphor laurel. Grime (1979)
indicates that plant dry matter production is influenced by a number of environmental
constraints which may include solar energy, moisture and nutrients. For camphor
laurel these have been identified as the influence of herbivory and predation of seed,
water-logging, salt tolerance and fire.

Absence of Significant Predation or Disease


There appears to be an absence of significant insect pests, fungal diseases and
epiphytic growth on leaves and to some extent the bark of mature trees although
several organisms have been noted to browse or infect the tissues of local camphor
laurel trees. Firth (1979) observed that camphor laurel has very few major predators in
the region which is often the case with an introduced plant outside its natural range.
This was also found by Scanlon et al. (2001). Firth (1979) also suggested that
camphor laurel may have efficient defense mechanisms such as inhibitory volatiles,
which have been used as an insecticide by humans.

However, seed predation and browsing of seedlings have been noted to be high due to
foraging wallabies and domestic livestock, with many seedlings not surviving the first
two years (Stewart 2000). Stewart (1995) found that seed predation, herbivory and
disease were major factors in reducing abundance of seedlings but not mature trees.
Limited numbers of seed were also identified in the seed bank. In Mullumbimby,
rodents were a major contributing browser of camphor seed, with 100% predation of
baits overnight. Stewart's (2000) further work also revealed considerable browsing of
seedlings by wallabies.

Livestock such as cattle and horses also browse young

camphor laurel saplings, stripping the young foliage, and this appears to contribute to
a reduction of recruiting trees in intensively grazed landscapes (Firth 1979). There
also appears to be limited insect predation of seeds under trees observed in northeastern NSW (author pers. obs.). Liu et al. (2002) identified a ribosome-inactivating
protein (RIP), cinnamomin, in the seed of camphor laurel. This was indicated to be a
28

type II RIP which has a carbohydrate binding activity and a bisulphide bond which
belongs in the Ricin groups of compounds, known to be extremely toxic. It was found
to act as a novel protein storage which liberates carbon, nitrogen and sulphur during
germination. However, it was also discussed that this is a toxic protein that acts on
eukaryotic and prokaryotic ribosomes and many RIPs are active as chemical defences
against viruses, fungi and insects. Cinnamomin in this case has also been
demonstrated to be toxic to insects such as the larvae of the mosquito (Culex pipines
pallens) and the cotton bollworm (Helico-verpa armigera) (Zhou et al. 2000). Levels
of this protein were found by Liu et al. (2002) to be high in camphor laurel seed,
accounting for 11% of total proteins. This is higher than many other storage proteins
in other plants such as soybean storage proteins (5-10%) (Liu et al. 2002). This toxic
protein may be the reason for the lack of observed predation of seed by insects.
Vertebrates such as rats appear to be less susceptible to Cinnamomin. Work by Neilan
(2004) found that rat predation of seed is substantial in the Mullumbimby area.

In Australia, Common and Waterhouse (1972) have identified that several species of
butterfly larvae feed on the leaves of camphor laurel along the east coast of Australia.
These include Chaetocneme beata, Graphium macleayanum, Graphium serpedon and
Polyura pyrrhus. Firth (1979) noted that the only insect pests in north-eastern NSW
included the butterfly Graphium serpedon charedon (C. and R. Felder) which feeds
on foliage, and Monalepta australis, which feeds on the flowers with both inflicting
little damage.

A few fungi and epiphylls (leaf micro-lichens) have also been observed on camphor
laurel trees and leaves (Firth 1979). Firth found that only one species of epiphyll, the
lichen Strigula elegans (Fee) Mull. Arg., results in some grey spotting of leaves. The
leaf fungus Pestalotiopsis sp. causes brown spotting and veinal browning. It has also
been noted in north-eastern NSW that a fungus, possibly Phytopthora cinnamomi, has
caused some damage and is sometimes apparent when physical damage such as
ringbarking from vines or fences has damaged the mature tree (Scanlon et al. (2001).
However, Firth (1979) identified that this fungus was more apparent on camphor
laurel trees growing in low lying or alluvial floodplains subjected to frequent waterlogging. The disease also becomes apparent when earth works occur near the tree

29

which damages the roots or trunk, possibly allowing entry of the pathogen as a result
of physical damage (author, pers. obs.).

Scanlon et al. (2001) identified a few species which are capable of inflicting some
damage on mature camphor laurel trees in Australia. These include the mountain
brushtail possum which consumes some leaves, shoots and green bark; rodent
predation of green bark; several species of butterflies which lay eggs on the leaves;
and several bird species including the brush turkey (Woodford 1993), king parrot
(Holmes 1987) and white headed pigeon (Firth 1979) which consume and destroy the
seeds.

Scanlon et al. (2001) also discusses a number of natural predators of camphor laurel
present in other countries. These include a nuclear polyhedrosis virus (Eriogyna
pyretorum), the boring and feeding larvae of Thymiatris sp., the camphor laurel leaf
caterpillar, (Charaxes bernardus fabricius), the larvae of Actias selene ningpoana in
Taiwan and the insect (Fruhstorferiola tonkinaris). In Florida, Firth (1979) also found
that camphor laurel thrips, (Crypotothrips floridensis Watson), and the large black and
white weevil, (Helipus apiatus Oliv.) do inflict some damage on the tree.

The observed low insect browsing, epiphyte and fungal development on leaves may
also result from the presence of a waxy leaf cuticle, allelopathic release of volatiles
from leaves, and possibly rapid drainage of the leaf resulting from the presence of a
drip-tip and relatively small leaf size, features which would aid the rapid drying of the
leaf surface (author, pers. obs.).

Survival in Marginal Habitats


Although camphor laurel grows vigorously in riparian locations it does not appear to
be common in seasonally or permanently wet locations, such as swamps and soaks.
Camphor laurel has been observed to germinate and survive as a scant slow-growing
tree of little competitive ability in fresh-water wetlands of Melaleuca quinquenervia
along coastal environments and in littoral rainforest gaps. It is able to achieve this
through its ability to germinate and persist on higher ground such as at the bases of
trees which form plinths or upraised platforms resulting from root binding of the soil
below the tree (author, pers. obs.). However, Firth (1979) indicates that the presence
30

of waterlogged soils results in a higher incidence of disease, assumed to be


Phytophthora, which he suggests decreases its competitive ability.

Camphor laurel does not appear to be present in saline environments such as estuaries
and mangrove forests of the region to any great extent (author, pers. obs.). Research
in China by Zhang et al. (2002) found that camphor laurel has a seedling survival rate
of 10% under saline conditions of 0.20 0.46% salt, and up to 80% under saline
conditions of 0.10 - 0.25% salt indicating a severe intolerance to salinity.
Tolerance of Soil Conditions
The work by Firth (1979) found that camphor laurel grew most prolifically on the
Ferrosol (Krasnozem) soils in the local region but also occurred on a variety of other
soils on both upper slopes and valley bottoms. Troup (1921) and Floridata (2004)
identify that the tree prefers well-drained fertile sandy soil and will not tolerate waterlogging. On poor soil Troup (1921) suggests that growth is usually stunted but it will
tolerate poor lateritic soils of good drainage. The tree can tolerate a pH range of 4.3 to
8 (Floridata 2004) indicating a relatively wide tolerance. However it does demonstrate
intolerance to highly acidic conditions such as found in acid sulphate soils and heathlands, and high alkalinity.
Secondary Strategy
The secondary strategy of camphor laurel involves the combined influence of being
both a competitive and stress tolerant plant in application of the three ecological
strategies portrayed by Grime (1979). Grime (1979) suggests that the r-K continuum
is confined in its explanation of the secondary strategies in application to plants and
can be further understood using the C-S-R strategy. The previous work by MacArthur
and Wilson (1967) defined r-selection as organisms possessing a short life-span and
large reproductive effort while K-selection are organisms which demonstrate a long
life expectancy while the proportion of energy dedicated to reproduction is small.

It can be identified from examination of the mature stage in the life-cycle of camphor
laurel that it displays both r- and K- attributes in its ecology using the model of
MacArthur and Wilson (1967), and Pianka (1970). The tree falls somewhere between
these two extremes of the continuum, perhaps more towards K- selection than r-

31

selection as competitive ability of the tree is most obvious in its growth cycle,
particularly during the mid to late growth stages. Specifically, some r- attributes may
include high seed production, seed dormancy i.e. delayed asynchronous germination,
relatively small seed size, long dispersal distance and light demanding nature; some
K- attributes may be longevity and competitive ability of the mature tree. The rattributes however, appear to be dominant in the early stages of growth and K- is most
prominent at maturity or near maturity.

Grime (1979) furthered this concept for specific application to plants using a
combination of the competitive - stress tolerance - ruderal (C-S-R) nature of the plant,
expressed during the mature phase of the growth cycle. When these principles are
applied to camphor laurel the following strategy emerges, identifying the tree as a
stress-tolerant competitor (S-C), as the tree exhibits several intermediate features in
its life-cycle which could place it in this category. The tree however, does not entirely
match this strategy either. Grime (1979) identified that a large number of trees and
shrubs occurring in unproductive environments or those that are persistent in the later
stages of vegetation succession in more fertile habitats, are S-C strategists. Other
attributes of this strategy also include intermediate longevity, fluctuating but
continuous seed production, and the ability to remain photosynthetically active
throughout the year (Grime 1979). However, in the case of camphor laurel, the
intermediate longevity appears to be replaced with ability to survive for very long
periods under highly competitive conditions in a mature phase forest (Firth 1979,
Valder 1999, Pakenham 2003). The tree also has an ability to produce large numbers
of short-lived seeds, which are widely dispersed and of a relatively small size. These
germinate and recruit mainly in open and disturbed habitats (Firth 1979, Panetta 2001,
Stewart 1995). These seed characteristics could also be viewed as being components
of an R strategy only occurring during the early growth stages and not in the
established phase of the plant. Grime (1979) also cautioned the use of this S-C
classification as the strategy had been little researched at that time. Camphor laurel in
this case, may be an anomaly, as a combination of S-C strategy occurs in its latter
growth stage but with a high longevity, and the additional features of an R strategist
early in its life cycle; displayed through its seed production and the requirement for
open, less competitive habitats for recruitment. Grime (1979) also suggested that the
concept of r- and K- selection proposed by MacArthur and Wilson (1967) and Pianka
32

(1970), only partly fits the model of the C, S and R strategy in plants. K- selection
was indicated by Grime (1979) to be consistent with the S plant strategy as these are
organisms that have a long life-span with a limited energy and resource allocation for
reproduction; and r- selection is similar to the R strategy where the life-cycle is short
and large amounts of energy are allocated to reproduction. The C strategist is
indicated to fall between the extremes of r- and K- selection. However S is not
recognized in the r-K continuum and was identified by Grime (1979) as a distinct
strategy which has evolved in situations where habitats are unproductive or where
severe resource depletion occurs as a result of the vegetation itself. Grime (1979)
further demonstrated that in order to identify the equilibria which exist between stress,
disturbance and competition in plants it is necessary to use a triangular model which
can be used to demonstrate the compromise between these factors in individual plants
or groups of plants of a similar strategy.

Regeneration
The regenerative strategies in camphor laurel involve efficient and prolific seed
production, dispersal and a specialized germination syndrome combined with
epicormic sprouting from branches, trunk, basal lignotuber and lateral roots.
Epicormic sprouting has been discussed previously and the following appraisal
reviews the flowering and fruiting syndrome in some detail:

Large numbers of seed are produced by this species (Firth 1979, Stewart 1995), and a
tree of 15-20 years or age in the Richmond-Tweed area produces around 113,000
fruits (Firth 1981). Firth (1981) found that fruits ripen in April-May and Stewart
(2000) identified that a lengthy seed rain occurs from March to September with a peak
in June, but this was variable in quantity and time from year to year. Firth (1979) also
identified that as the tree develops its canopy seed production increases over time.
This seed production may exceed the quantified numbers established by Firth (1979)
for younger trees.

Pollination of the tree is thus very efficient due to such high seed numbers are
produced. Firth (1979) identified that pollination may result from the presence of two
Dipteran flies, Parapeleosepsis plebia (del Meij) and Desmometapa ciliata Hendel,
and through self-pollination, with apomixis found to be ineffectual. This suggested
33

that both cross-pollination and self-pollination occur. Firth (1979) however was
uncertain of the pollination role played by the possible Dipteran pollinators.

Productivity of the tree at the time of seed production and its relationship to moisture
availability is also an important factor, as the large volume of seed produced would
require an efficient sugar production. The specialized leaf abscission behaviour noted
by Addicott (1978) may play a role in maximizing the production of sugars at the time
of fruit set. The requirement for high sugar levels during fruit development also seems
to be an important factor when it is considered that the seed of camphor laurel
contains a protein, cinnamomin in high concentration which provides nitrogen,
sulphur and carbon during germination (Liu et al. 2002) requiring a high demand for
sugar production. Carbohydrate production therefore would seem to be a supportive
factor in producing such large numbers of seed which would require high sugar levels
during protein synthesis. This would also mean that sugar production and fruit set are
closely associated with environmental factors which influence growth such as
moisture, temperature, solar radiation variation and the resultant influences on
hormone activity within the plant. It is most likely that the higher temperatures and
moisture levels experienced in north-eastern New South Wales at the time of vernal
abscission (growth) and germination as compared with China where moisture at this
time is absent, has provided an opportunity for the tree to proliferate in the region.
This was identified by Chen et al. (2004) as being the reason for the rarity of the tree
in China i.e. an absence of moisture during flowering, fruiting and germination.

Camphor laurel demonstrates a highly efficient seed production and a widespread


dispersal. This occurs due to a number of native birds (Firth 1979, Stewart 1995,
Holmes 1987, Date et al. 1991, Neilan 2004), fruit bats including the black flying fox
(Holmes 1987), by water movement in creeks and rivers with seeds remaining viable
for up to forty days underwater, and through rain-wash below trees and in steep
terrain (Firth 1979). The bird dispersers are varied and include many birds from
various feeding guilds (Neilan 2004). Firth (1979) estimated that dispersal distance
can be as great as 60 km for bird dispersal and perhaps as far as 108km for water
dispersal in creeks and rivers. Such efficient dispersal over long distances does
provide for the rapid advancement of the species over a broad range of geographical
locations and vegetation types, as has been demonstrated by Neilan (2004) for bird
34

dispersal. This study on rainforest regeneration in the Big Scrub region compared the
number of frugivorous birds and the diversity of the vegetation in camphor laurel
dominated forests located at various distances from the largest source area for seed in
the region (the Nightcap Range). A relationship between distance from seed source
and diversity of bird dispersed plant species in camphor laurel stands was identified in
this study. Diversity of bird dispersed species recruiting in the camphor laurel forests
was found to decrease with distance from the source area and suggested that stands of
this tree support a diverse assemblage of avian frugivores numbering twenty seven
species in nine frugivorous guilds. Neilan (2004) observed these frugivores in
camphor laurel forest but did not note whether they were feeding on camphor laurel
fruit. Table 3 lists Neilan's (2004) frugivores and compares them against the known
camphor laurel seed feeders listed in the literature, providing a list of fourteen known
camphor laurel frugivores, of which nine were recorded by Neilan (2004).
Several characteristics of camphor laurel seed provide for a specialized germination
strategy. Camphor laurel has a reliance on the potential for germination with a
delayed asynchronous germination rather than a persistent seed bank (Panetta 2001).
In early work by Firth (1979) it was suggested that seeds may remain viable in the soil
for perhaps three years providing for a persistent seed bank. In contrast, Ghosh (1977)
in India found that seeds of this species remain viable for a period of 200-259 days.
Interestingly, Panetta (2001) during field trials in southern Queensland found that
viability is also lost quickly under field conditions with only 1% of seed remaining
viable after 12 months. This he identified is similar to a few other woody weeds such
as *C. sinensis, *S. terebinthifolius and *Ligustrum spp. which also have transient
seed banks. The syndrome of regular and prolific seed production of limited seed
longevity is a common feature of exotic species that have invaded moist natural
ecosystems in subtropical Queensland. This means that the reliance is placed more on
the potential for recolonization rather than on an ability to have a persistent seed bank
(Panetta 2001).

Stewarts (2000) study also demonstrated that seed germination occurred from
November to July but was also variable in both time and quantity from year to year.
An explanation for the reason why germination was so varied was obscure, being
identified as possibly due to variation in fruit production or the abundance and

35

Table 3. Comparison of frugivores identified as present in the camphor laurel forests by


Neilan (2004) using the feeding guild structure applied in that study, compared with
known dispersers of camphor laurel seed cited in the literature.
Guild and Species (Neilan 2004)

Research Identifying Species as Camphor


Laurel Seed Feeder

Large-gaped major:
Channell-Billed Cuckoo
Figbird
Topknot Pigeon
Wompoo Pigeon

(Firth 1979)
(Firth 1979, Date et al. 1991, Scanlon et.al. 2001)
(Date et al. 1991, Scanlon et.al. 2001)

Medium-gaped major:
Rose-Crowned Fruit Dove

(Holmes 1987, Date et al. 1991)

Large-gaped mixed diet:


Green Catbird
Paradise Riflebird
Pied Currawong
(Firth 1979)
Satin Bowerbird
Medium-gaped mixed diet:
Lewin's Honeyeater
(Holmes 1987)
Small-gaped major:
Mistletoebird
Small-gaped mixed diet:
Silvereye
Varied Triller
Minor Frugivores:
Australian Magpie
Grey Butcherbird
Torresian Crow
Scarlet Honeyeater
Seed Grinders:
Australian Brush Turkey
Australian King-Parrot
Brown Cuckoo-Dove
Crimson Rosella
Eastern Rosella
Rainbow Lorikeet
Sulphur Crested Cockatoo
White-Headed Pigeon
Wonga Pigeon

(Woodford 1993)
(Holmes 1987)

(Firth 1979, Date et al. 1991)

Other Species Recorded as Dispersers of Camphor Laurel Seed


Superb pigeon
Olive-backed oriole
Noisy miner
Scaly-breasted lorikeet
Black-faced cuckoo shrike

(Scanlon et al. 2001)


(Firth 1979)
(Firth 1979)
(Holmes 1987)
(Holmes 1987)

36

movement of dispersers such as rainforest pigeons. Chen et al. (2004) however, in


studies in China, identified that the trees rarity there as compared with those
countries in which it has become an environmental weed was due to its fruiting period
and germination occurring from October to December, during the dry season which
extends from October to March in Southern China. This reduction in moisture during
the fruit development and germination period was found to reduce the number of
seeds produced and the likelihood of seed successfully germinating, thereby
contributing to the rarity of the tree in China (Chen et al. 2004). In comparison,
Stewarts (2000) peak seed rain period at Mullumbimby was from April to September
(a winter dispersal) during a period of reducing moisture and temperature, while seed
germination occurred from November to July (a period of higher initial moisture
availability and temperature). However, Stewart's (2000) work near Mullumbimby
also recorded a peak seed rain density of 1-3 seeds per m and a seedling emergence
occurring between November and July following the winter seed rain, taking one
month to commence germination. This field recording was identified as being
consistent with Firths (1979) laboratory trial which demonstrated a delayed
asynchronous germination. The seed emergence syndrome of a delay, followed by
variability in seedling emergence over time was discussed by Stewart (2000) as being
a result of inherent dormancy characteristics of the seeds. Firth's (1979) earlier work
also found that this delayed germination varies from between 4-20 weeks under
laboratory conditions, supporting Stewarts view that germination delay is due to
genetic variation. Stewart (2000) indicated that this strategy of delayed germination
enables early germination during the wet season and therefore a maximum growing
period for camphor laurel, providing a potential competitive advantage. However, the
work by Chen et al. (2004) in China clearly demonstrates that seed germination is
limited by the moisture regime and therefore Stewarts variability in germination may
also result from moisture variation in the soil from month to month, in addition to the
identified genetic variation.

In a study of moisture content and temperature influences on seed germination and


storage of camphor laurel seed, Chien and Lin (1999) found that the seeds are
recalcitrant but not typically so. Recalcitrant seeds are both sensitive to moisture loss
and low temperature, as opposed to species that have an orthodox storage behaviour,
drying to a low moisture content (<10%) with limited effect from low temperature.
37

They found that the viability of camphor laurel seed was lost entirely following an 8
month period at temperatures of 5C and 15C, where moisture content was higher
than 10.4 %. Under field conditions in Taiwan, the moisture content of fresh seed was
28.6%. However, when moisture content was lower than 6.7 % at these temperatures,
60% germination occurred after 12 months and at lower temperatures of -20C, seed
rapidly lost viability after only one month. When seed was at a moisture content of
21.1% and a temperature of 15C, germination increased to 50% after one month and
viability was maintained for only 4 months. Chien and Lin (1999) further add that
camphor laurel seeds can tolerate desiccation to the same extent as orthodox seed but
when temperatures are extremely low they are more sensitive to damage. In seed
studies in China by Chen et al. (2004) fresh fruit was found to have 100% viability
when moisture content was at 46.3%. Seed retained viability in an open air situation
for one month when moisture content decreased to 25% but, over a two month period,
viability was found to fall to 70% with a moisture content of 18%.

In Australia, Firth (1979) and Anon. (1998) found that the germination of camphor
laurel is highly dependent on lengthy periods of soil moisture retention and that
surface seed exposure to desiccation results in limited germination viability. In a
further experiment by Panetta (2001) moisture availability was further identified as an
important factor in seed germination in southern Queensland. It was found that
surface sown seed had virtually no germination under natural rainfall while irrigated
seed had almost 100% germination. Panetta indicated that surface lying seeds will not
germinate unless in micro-sites which provide protection and an adequate surface
retention of moisture. The experiment provided a highly significant interaction
between the water regime and the burial status on emergence of the seed i.e. an
obligate dependence on burial. However, the work also found that the highest
periods of germination occur when mean weekly temperatures were 22 C. An
observation on the potential for autumn to late winter germination directly following
seeding revealed an absence of germination in the field.

The findings by Firth (1979), Chien and Lin (1999), Anon. (1998) and Panetta (2001)
clearly identified that low moisture content, low temperature and soil surface
placement of the seed as important interrelated and limiting factors in seed viability,
placing limitations on germination in situations of low moisture and temperature.
38

In an experiment by Russell (1919) in Florida, USA, removal of the seed pericarp of


camphor laurel stimulated germination but this was found to vary from month to
month. The germination ranged from as high as a 51% increase to as low as 10%
when compared without seed without pericarp removal. Firth (1979) also found that
pericarp removal from the seed resulted in an increase in germination percentage, but
this too was variable on a monthly basis ranging between 30% to as low as 2% when
compared with seed with the pericarp intact. Ingested seed by birds was also
demonstrated to dramatically increase germination relative to seeds with flesh intact
or flesh removed. Similarly, Panetta (2001) identified that the retention of the pericarp
at planting of camphor laurel has a significant influence on reducing germination to
less than 6% and was slower as a result of this inhibition. This identified a significant
dependence on frugivorous removal of the pericarp to enhance germination in the
natural environment. Stewart (2000) found differences in germination due to pericarp
removal by various dispersers such as the Topknot pigeon (Lopholaimus antarctica)
which pass the seed, and species such as the Pied currawong (Stepera graculina)
which regurgitate the seed with flesh intact. Anon. (1998) further added that removal
of the pericarp followed by immediate burial increased seed germination from 10%
after 52 weeks for peeled and stored seed to 56% after 20 weeks for peeled and
immediately planted seed.

In work involving an attempt to stimulate the germination of camphor laurel seed by


Bahuguna et al. (1987) in India to overcome its erratic and slow germination for
forestry purposes, it was found that puncturing of the seed coat following removal of
the pericarp resulted in no stimulation within 10 days. Partial removal of the seed coat
resulted in 9% germination; and full removal produced 100% germination within that
period. This suggested that the seed coat itself contains the inhibitory mechanism.
Bahuguna et al. (1987) however, attributed this to the mechanical resistance of the
seed coat and the inability of the embryo to rupture it, but did not refer to any
possibility of inhibitory compounds being present. However, Firth (1979) in northeastern NSW found that seed coat removal provided no advantage, indicating that
dormancy could also result from inhibitory compounds elsewhere in the seed.

Camphor laurel has been observed to appear most frequently in disturbed forest and
open country which receives relatively high light levels (Firth 1979, Stewart 1995,
39

Schenk and Wallace 1996). Little seed germination has been noted in forests which
have minimal recent disturbance and high canopy closure (author, pers. obs.). In these
forests camphor laurel germination has been observed by the author to be confined to
large gaps of high light intensity or as scattered individuals of low vigor under a
closed canopy. This may identify camphor as a 'large gap' colonizer suited to high
light levels resulting from disturbance i.e. a pioneer. Ecological advantage is gained
for the tree through recruitment into an open disturbed ecosystem where there is less
competition for light and resources and a greater chance of an available niche (author,
pers. obs.). This observation is supported by work in China by Chen et al. (2004) who
demonstrated a considerable dormancy period of 55 days under light conditions (25
mol/m2 per s over 12 hour day) and 69 days under dark conditions (total darkness),
which lasted for more than 3 months. The percentage of germination in the study was
also found to be low being only 16% in the light condition and 2% in the dark
condition with germination commencing 2 weeks later than seeds in the light
condition. In the forest situation however, seed was found to germinate after 3 months
with low germination of 1.6%.

Firth (1979) found camphor laurel seed germination in a variety of open habitats such
as fence-lines, regenerating forests and below perch sites. Dunphy (1988) indicated
that camphor laurel was intolerant of high shade conditions and germination and
survival occurred at the highest levels near edges of remnants, around single trees, and
in regrowth and degraded farmland. Stewart (1995, 2000) also identified seed rain
around single perching trees in open areas. Interestingly, a high and low response to
light was identified for camphor laurel seed germination by Cornelissen et al. (1994)
who discovered in China that seedlings of camphor laurel reach maximum size under
an intermediate light regime of 33-55% full light, identifying that both low light of
18% and high light of 100% were less favourable for germination and seedling
development. However this work also found that independent of the light regime in
the forest, the seedlings of the tree attained a higher biomass accumulation, stem
height, stem diameter, total leaf area and number of leaves than the other five tree
species growing in association with it. In work in north-eastern New South Wales by
Stewart (1995, 2000), low survival of camphor laurel seedlings below a dense
camphor laurel canopy was attributed to low light levels. Stewart further noted that
seedling emergence under the canopy approached zero within two years of seed rain.
40

Factors which may explain this were identified as low light levels, herbivory, and
disease. The conclusion drawn by Stewart (2000) was that camphor laurel will not
invade sites which possess a full canopy whether it be advanced regeneration or
mature forest cover. In support of this, Diana et al. (1997) in Japan found that
camphor laurel seedlings exposed to low levels of red and far-red light as would be
expected below a tree canopy, had reduced branching number, stem diameter and root
weight with three to six times higher stem elongation, higher stem height and lower
specific leaf area.

Chen et al (2004) in further glasshouse trials in China found that greater leaf growth
of seedlings occurred under higher light intensities of 54.6 mol/m2 per s having a
very significant (P< 0.01) effect on leaf number. Reducing light intensity of 36.4 and
18.2 mol/m2 per s resulted in decreased leaf number. However, light levels had no
significant effect on shoot height; although seedlings appeared taller under lower light
levels of 18.2 mol/m2 per s. Higher light levels also produced a significant effect on
branching, producing earlier and higher numbers of branches (Chen et al. 2004).

1.2.4 Summary of Ecological Characteristics


Grime (1979) produced a table which identifies the attributes of the competitive,
stress tolerant and ruderal plant strategies using eighteen distinct features.
Application of this table to camphor laurel identify sixteen competitive characteristics
and ten stress-tolerant characteristics which identifies the tree as being a Competitive
- Stress-tolerant strategist. The sixteen competitive and ten stress-tolerant
characteristics of camphor laurel are presented in Table 4.

Grime (1979) also identified several morphogenetic responses to desiccation, shade,


and nutrient stress in the competitive, stress-tolerant and ruderal plants in three
varying habitats; these included early successional stages of productive and
undisturbed habitats, continuously unproductive habitats or late stages of succession
in productive habitats, and severely disturbed and potentially productive habitats.
The composite nature of the Stress-tolerant competitor (S-C) strategy of camphor
laurel and its ability to be a successful invasive and persistent plant is emphasized by
these morphogenetic response groups.

In application of these principles of

morphogenetic response listed by Grime (1979) a most interesting pattern emerges.


41

Where there is early succession in productive and undisturbed habitats and stresses
are mainly plant induced, the response is to maintain high absorption of water and
nutrients to ensure dry matter production under stress, enabling success in
competition. In continuously unproductive habitats where stress is relatively constant
due to climate and/or soil, or in the later stages of succession in productive habitats

Table 4. Summary of the competitive and stress-tolerant attributes applied to

Camphor laurel following Grimes C-S-R strategy (1979).


Competitive Attributes:

Life-form as a tree
Leaf canopy high and dense, extensive lateral spread of canopy and roots
Exceptional longevity 700 to 3000 years?
Short lived leaves biannual leaf shedding
Leaf production coincides with periods of maximum potential productivity
Flowers produced prior to maximum potential productivity through vernal abscission
Trees flower biannually through vernal abscission
Amount of sugar production devoted to seed is small
Perennation through dormant silleptic buds in the leaf axles, epicormic buds associated with roots, lignotuber, trunk and
branches, and by seed
Regenerative strategy is seasonal regeneration in vegetation gaps and vegetative expansion
Growth rate is rapid
Response to stress rapid which maximizes vegetative growth
Photosynthetic response is strongly seasonal and highly efficient
Photosynthate and nutrients incorporated into vegetative structure with storage for growth in the next season
Leaf litter is produced in large quantity and is slow to decompose, releasing allelochemicals
Palatability to vertebrate herbivores is high in the early growth stage while invertebrate browsing is low

Stress-tolerant Attributes:

Life-form as a tree
Small leathery leaves displaying some sclerophylly
Long life-span 700 to 3000 years?
Evergreen leaf phenology
Small proportion of annual sugar production devoted to seed
Stress tolerant leaves and roots
Regeneration strategy towards vegetative expansion with seedlings being persistent
Opportunistic photosynthesis which is uncoupled from vegetative growth through vernal abscission
Ability to store photosynthate in stems and roots in the mature tree as well as seedlings
Palatatability to unspecialized herbivores particularly insects is low in the mature tree

the stress tolerant nature of the tree enables the conservative use of water, nutrients
and photosynthate providing persistence for long periods, during which dry-matter
42

production is limited. This appears to be consistent with the longevity of the camphor
laurel tree and its persistence in a mature forest situation. Interestingly, both the
competitive and stress tolerant strategies result in failure of seed production in
severely disturbed, potentially productive habitats such as under conditions of
drought, thereby reducing the chance of rehabilitation after disturbance. Camphor
laurel may be able to circumvent this by having a seed that is capable of surviving for
at least a year in the environment (Pannetta 2001), with germination dependant on
specific micro-site moisture content (Firth 1979).

These reviewed adaptations and strategies clearly demonstrate the complex


interrelationships that may exist in the life-cycle of camphor laurel both in
environmental, metabolic and ecological factors influencing its success or restraining
its growth. Clearly it identifies the enormous capacity of the tree to compete
successfully in a regenerating forest. Additionally, these discussed capabilities act as
a guide for further research and discussion on the ecological and adaptive strategies of
the tree.

1.3 Further Information Needs

1.3.1 Introduction
Although many adaptations and strategies may exist for camphor laurel which may
provide for ecological advantage in a regenerating plant community, a single topic has
been selected for this study.

In this research, allelopathy in camphor laurel is addressed with the intention of


contributing some answers to the perplexing and somewhat difficult question of how
camphor laurel is so successful, and how this component of its strategy may be
influencing plant communities. This reputed allelopathic effect on the regenerating
plant assemblage has been based, in many cases, on visual observation of species
presence and abundance, and the visual impact of the plants biomass. Interestingly,
little research has been undertaken to assess whether camphor laurel allelopathy is an
important contributing factor in the success of this species, and most importantly,

43

whether allelopathy is exhibited at a significant level in the plant assemblage as a


whole or whether succession in native Australian vegetation may be altered.
Overall, research into the allelopathic influence of camphor laurel on the plant
communities of the region placed into the overall framework of the trees ecology
should be able to contribute to answering the following fundamental questions:

How is camphor laurel able to be such an efficient and effective intrusive plant
i.e. what physical and ecological factors are involved?

Is the intrusive nature of camphor laurel in regenerating native forests altering


plant succession and, if this is of consequence, could this be a threatening
process to some plant assemblages or species? and

What might be the implications for management including regeneration


techniques in camphor laurel-invaded plant communities following the
provision of further analyses of the trees allelopathy within the trees overall
ecology?

1.3.2 Definition of Allelopathy


The term allelopathy is referred to by Molisch (1937) and Rice (1984) as being:

any direct or indirect harmful or beneficial effect by one plant (including microorganisms) on another through production of chemical compounds that escape into
the environment.

Importantly, the chemical process of allelopathy may produce both positive and
negative growth responses (Muller 1971, Rice 1984, Rizvi et al. 1992) and can be
considered to be a biochemical interaction among plants (Rizvi et al. 1992).

These biochemical interactions are distinct from plant competition which involves the
removal or reduction of an environmental factor required by another plant occupying
the same habitat (Grime 1979, Rice 1984). Collectively however, allelopathy and
competition are referred to as interference studies (Rice 1984).

44

Grime (1979) indicated that allelopathic compounds are secondary plant metabolites
often having no distinct function. However it is cautioned that many compounds such
as terpenoids and alkaloids often exist in a state of dynamic equilibrium within plants.
These compounds may not be just end products of metabolism having no function and
may have yet unknown reasons for being present within the plant. Such compounds
may not have evolved merely to prevent herbivory or act as allelopathic compounds
(Grime 1979). Interestingly, Grime identified that allelopathy, which can contribute to
dominance of a species in a plant community, is most commonly expressed in the
competitive-ruderal group.

Whittaker (1970) indicates that the chemicals involved are found in almost all plants
and the effects can be obvious in plant communities. The plant parts involved may be
above or below the ground surface and allelopathy can occur through the process of
leaching, volatilization, exudation, excretion and decay either with or without the aid
of micro-organisms (Whittaker 1970). The allelochemicals have been discussed in
detail by Rice (1984) and are considered to be mostly secondary metabolites (Rizvi et
al. 1992). These may include: phenyl propanes, acetigens, terpenoids, steroids and
alkaloids (Whittaker and Feeny 1971). Rice (1984) however further identified these
allelochemical groupings: simple water soluble organic acids, straight chain alcohols,
aliphatic aldehydes and ketones; simple unsaturated lactones; long-chain fatty acids
and polyactetylenes; naphthoquinones, anthroquinones and complex quinines; simple
phenols, benzoic acid and derivatives; cinnamic acid and derivatives; terpenoids and
steroids; amino acids and polypeptides; alkaloids and cyanohydrins; sulphides and
glucosides; and purines and nucleosides (Figure 1). Rice (1984) demonstrated in
Figure 1 that inhibitors arise either through the acetate or the shikimic acid pathway
with several types of inhibitors, which originate from amino acids, being produced
through the acetate pathway. According to Niesh (1964), Whittaker and Feeny (1971),
and Robinson (1983) these may include some of the amino acids, polypeptides,
alkaloids, and sulphides, as well as the purines and nucleosides. Apparently some of
the other amino acids also arise from phenylalanine or tyrosine which is formed from
shikimic acid.

The chemicals may operate singly or in combination with others to produce the
allelopathic effect which is also dependant on the quantity and time of exposure
45

(Asplund 1969, Rice 1984, Cheng 1992). The effect of these chemicals can also be
intensified by external environmental factors such as shade, soil drought, nutrient
levels, root competition and animal influences (Whittaker 1970). Rizvi et al. (1992)
indicate that they can also be indirect or direct in their operation. In an indirect way
the soil itself may be altered through structural change, nutrient availability and
influence beneficial or harmful micro-organisms and invertebrates resulting in an
hydrolyzable tannins
digallic acid, etc

carbohydrate

gallic acid and protocatechuic acid


pyruvate
naphthoquinones and anthraquinones
dehydroshikimic
acid
amino acids and polypeptides
alkaloids and cyanohydrins
shikimic acid
sulphides and mustard oil glycosides
amino acids

purines and nucleosides


simple phenols, benzoic acid and
derivatives

acetate

cinnamic acid
derivatives
mevalonic
acid

courmarins
flavonoids

condensed tannins

terpenoids
and

steroids

water-soluble organic acids, straight chain alcohols,


aliphatic aldehydes, and ketones
simple unsaturated lactones
long chain fatty acids and polyactetylenes
naphthoquinones and complex quinines
phloroglucinol and polyphloroglucinols

Source: after Rice 1984

Figure 1. Biosynthetic pathways identified as probable origins of allelochemic agents.


Source: after Rice 1984

alteration to plant population (Rizvi et al. 1992). With the direct effect, Rizvi et al.
(1992) identify that the allelochemicals influence plant structure and metabolism

46

through changes to cytology and ultrastructure; the balance of phytohormones;


membranes and their permeability; germination of pollens and spores; the uptake of
minerals; stomatal response, pigment synthesis and photosynthesis; respiration;
protein production; leghaemoglobin synthesis and nitrogen fixation; specific enzyme
activity; conducting tissue; water relations of plants, and genetic material. Buchholtz
(1971) suggests that mineral nutrition of the plant may be strongly affected through
reducing available soil nutrients, root absorptive area, capacity to absorb nutrients,
ion-transfer in root membranes and transport of nutrients to shoots.

These direct and indirect allelopathic effects influence plant growth and hence
community structure and composition in several ways. The sequence of succession
may be altered by reducing the time taken to replace a species by a successor,
suppression or reduction of growth of the surrounding species delaying or preventing
replacement, inhibition of soil organisms and changes to decay processes (Whittaker
1970).

1.3.3 Allelopathy and Camphor Laurel


Early work by Muller (1966) suggested that hydrophobic terpenoids such as -pinene,
cineole, and camphor may be present in the leaves of *C. camphora and through the
process of volatilization under drought conditions could be released to the
environment. In the work by Asplund (1969) the phytotoxic monoterpenes such as
camphor, pulegone and borneol applied to the germinating seed of radish (*Raphanus
sativus), wheat (*Triticum aestivum) and oats (*Avena sativa) as individual
compounds and in combination found that a marked synergistic allelopathic effect
developed where these were applied in combination. Chou et al. (1989) on the other
hand in an experiment in Taiwan investigating the phytotoxic effects of kikuyu (*P.
clandestinum) and various tree species such as *C. camphora, *Alnus formosana and
*Zelkova formosana on the germination of vegetables and pasture grasses found that
as the aqueous extract percentage of *C. camphora increased, radicle growth
decreased for perennial rye (*Lolium perenne), ryegrass (*Lolium multiflorum) and
Chinese cabbage (*Brassica oleracea). The papers by Asplund (1969) and Chou et al.
(1989) suggest that the allelopathic effect in *C. camphora originates from the
phenolics and the synergistic combination of the monoterpenes. Of note, Asplund
(1969) and Whittaker (1970) indentify that the terpenes are responsible for a strong
47

inhibitory response to germination as well as to seedling growth through inhibiting


respiration and increasing the plants susceptibility to environmental stress during dry
periods. Furthermore, Whittaker (1970) indicates that the phenols contribute a
protective function to plants against fungi, bacteria and viruses and discourage
browsing by animals due to their bitter taste.
Muller et al. (1964), Muller (1974) and Rice (1984) suggest that many volatile
compounds may either be lost to the atmosphere or adsorbed on to soil particles.
Interestingly, Muller (1966) indicated that the hydrophobic terpenoids present in
camphor laurel, such as -pinene, cineole and camphor, could be released by
volatilization from leaf litter during periods of drought. Firth (1979) recognized that
soil release of aromatic compounds was occurring, as a distinctive odour was found to
be present in soil samples taken from the vicinity of camphor laurel trees, although no
effect on the growth of plants in this soil resulted in a glasshouse trial. The nature of
allelopathy as indicated by Rice (1984), not only involves a complex and varied group
of biochemical compounds and pathways, but is also influenced by variation in effect
due to their concentration, influences due to varying environmental and soil
conditions, and variability in species response to the compounds. It is possible that
any effect observed due to allelopathy in camphor laurel would not only be due to the
oils themselves but due to a complex set of these interacting factors including the
possible presence of other biochemical compounds. However, inhibitory allelopathy
has been indicated by Rice (1964, 1965 and 1984), Muller et al. (1964), Whittaker
(1970) and Muller (1966, 1974) for the essential oils cineole and camphor, as well as
many of the other lesser terpenoid oils found in this tree. The allelopathic pathways
identified by Rice (1984) in Figure 1 demonstrate the possibility for many of these
compounds to be present, as the pathway to terpenoid production through pyruvate,
acetate, and mevalonic acid may enable a range of other potential allelochemics to be
present.

Firth (1979) discussed the possibility of allelopathy in the Australian camphor laurel
trees noting the observation by farmers and gardeners that adjacent plants to mature
camphor laurels trees were being poisoned. A trial on the effect of allelopathy on
seed germination by Firth (1979) was mostly inconclusive being identified as a result
of the use of soil low in volatiles, and limited exposure time. However, in the second
experiment by Firth (1979) which used aqueous extracts from the roots and leaves of
48

the camphor tree, a significant difference in germination between the untreated and
treated seeds of *Plantago lanceolata, *Lolium rigidum and *Hypochoeris radicata
was identified. These three introduced plants had consistently lower germination
percentages, although there was no evidence of a reduction in vigour, shoot length,
leaf number and colour in the treated seedlings. However, Firth (1979) suggested that
under field conditions allelopathy may be more effective, as seeds may be exposed to
the extracts for longer periods than in the experiment which may result in the
impairment of growth or even induce mortality, as evident in studies of camphor
laurel seedlings under parent trees. Even though many of these phytotoxic compounds
are either volatilized rapidly into the environment or break down on exposure to soil,
regular replacement of these chemicals occurs through leaf fall and aqueous removal
by rain, fog drip or direct release from the plant (Rice 1984). The leaf extracts of the
camphor tree have also been identified as being significantly inhibitory on the
germination of *Pennisetum clandestinum, *Cedrela odorata, Toona ciliata (T.
australis), Casuarina cunninghamiana and its own seed by Johns (1994).

Hirota and Hiroi (1967) identified two chemo-types of camphor laurel in Queensland,
which were the camphor and cineole types assessed using the aroma of the leaves.
In work by Stubbs and Brushett (2001), Stubbs et al. (2004) identified a range of
terpenes and essential oils particularly cineole and camphor which give the chemotypes of *C. camphora present in Australia their name due to the dominating
abundance of these compounds. These varied terpenes occur in the leaves, branches
and roots of the tree (Stubbs and Brushett 2001; Stubbs and Specht 2004). However,
these studies found fourteen oils in the leaves but at varying concentrations. These
leaf oils included -pinene, camphene, sabinene, myrcene, -pinene, limonene, 1,8cineole, camphor, terpinen-4-ol, citronellol, -caryophyllene, germacrene, -selinene,
and allo-aromadendrene (Stubbs and Brushett 2001). Figure 2a and b show the
concentrations of these leaf oils, while Table 5a and b provide the measure of
consistency between the sampled trees for each of the oils. This measure, providing
both standard deviation and confidence interval, identifies a high level of consistency
between many of the oils within each chemo-type. Apart from the leaves, these oils
were also found to be in the roots, trunk, branches and fruits of the tree. In addition,
safrole and an isomer of santalene were also identified. These compounds were found
to vary according to chemo-type and physical location on the plant (Stubbs et al.
49

2004). The work of Chou et al. (1989) in Taiwan also found the presence of a number
of phtytotoxic phenolics in *C. camphora, which included: -resorcylic acid, (a).
80

Oil
components:
O il C o m po ne nt s :
A = -pinene
B = camphene
C = sabinene
D = myrcene
E = -pinene
F = limo nene
G = 1,8-cineo le
H = campho r
I = terpinen-4-o l
J = citro nello l
K = -caryo phyllene
L = germacrene
M = -selinene
N = allo -aro madendrene

70
1,8-cineole

% oil in leaf

60
50
40
sabinene
30
-pinene

citronellol

m yrcene

20
10
0
A

F
G H
I
oil components

s o urc e : adapted fro m Stubbs & B rushett (2001); pers. co m Stubbs 2003

(b).
80

Oil
components:
O il C o m po ne nt s :

camphor

A = -pinene
B = camphene
C = sabinene
D = myrcene
E = -pinene
F = limo nene
G = 1,8-cineo le
H = campho r
I = terpinen-4-o l
J = citro nello l
K = -caryo phyllene
L = germacrene
M = -selinene
N = allo -aro madendrene

70

%oil in leaf

60
50
40
30

limonene

-pinene

b-caryophyllene

20
10
0
A

F G H
I
J
oil com ponents

s o urc e : adapted fro m Stubbs & B rushett (2001).

Figure. 2 Camphor laurel leaf oil (terpenoid) content of the: (a) cineole;
(b) camphor chemo-types. Source: adapted from Stubbs & Brushett (2001).

50

Table. 5 Confidence Interval at 95% (0.05) and Standard Deviation of the oils of
camphor laurel in: (a) cineole chemo-type; (b) camphor chemo-type.
(a).
Oil Components

Mean

sd

CI

4.533
0
16.917
3.2
1.5
0.75
50.85
0.2
0
9.05
1.1
0.75
1.183
1.025

0.207
0
0.637
0.141
0.082
0.5
3.1
0.447
0
0.539
0.972
0.599
0.972
0.05

0.165
0
0.51
0.113
0.08
0.49
2.481
0.392
0
0.432
0.777
0.479
0.778
0.049

Sample size
A = -pinene
B = camphene
C = sabinene
D = myrcene
E = -pinene
F = limonene
G = 1,8-cineole
H = camphor
I = terpinen-4-ol
J = citronellol
K = -caryophyllene
L = germacrene
M = -selinene
N = allo-aromadendrene

6
6
6
6
4
4
6
5
6
6
6
6
6
6

(b).
Oil Components

Mean

sd

CI

4.475
2.48
1.015
1.495
1.855
3.51
1.665
68.26
1
1
3.75
2.115
2.6
1.605

0.183
0.154
0.037
0.083
0.119
0.251
0.743
3.85
0
0
1.813
1.192
1.527
0.635

0.08
0.068
0.016
0.036
0.052
0.11
0.326
1.687
0
0
0.795
0.522
0.669
0.278

Sample size
A = -pinene
B = camphene
C = sabinene
D = myrcene
E = -pinene
F = limonene
G = 1,8-cineole
H = camphor
I = terpinen-4-ol
J = citronellol
K = -caryophyllene
L = germacrene
M = -selinene
N = allo-aromadendrene

20
20
20
20
20
20
20
20
20
20
20
20
20
20

51

Figure 3. Quantitative comparison of the phytotoxic phenolics from the leaf of


camphor laurel. Source: adapted from Chou et al. (1989).

resorcylic acid, -hydroxyphenylacetic acid, protocatechuic acid, vanillic acid and coumaric acid (Figure 3). This study however, did not differentiate these compounds
at the chemo-type level so it is unknown which chemo-type of the tree these
phytotoxins represent or whether they are present in the Australian trees.

Muller et al. (1964), Muller (1974) and Rice (1984) suggest that many volatile
compounds may either be lost to the atmosphere or even be adsorbed on to soil
particles. Interestingly, Muller (1966) indicated that the hydrophobic terpenoids
present in camphor laurel, such as -pinene, cineole and camphor, could be released
by volatilization from leaf litter during periods of drought. The nature of allelopathy
as indicated by Rice (1984), not only involves a complex and varied group of
biochemical compounds and pathways, but is also influenced by variation in effect
due to their concentration, influences due to varying environmental and soil
conditions, and variability in species response to the compounds. It is possible that
any effect observed due to allelopathy in camphor laurel would not only be due to the
oils themselves but due to a complex set of these interacting factors including the
possible presence of other biochemical compounds.

52

However, camphor laurel allelopathy has not been fully assessed as an 'operating field
component' in the general ecology of the tree as a representative sample of species
occurring in or near camphor laurel trees has not been investigated.
No references are known relating to the influence of aqueous leaf leachates of
camphor laurel on soil algal populations and growth. However, Choi et al. (1999)
demonstrated that some bacteria and fungi are strongly influenced by the
allelochemicals found in *C. camphora while other studies by Abivardi (1979),
Tiwari and Dixit (1994), Chao et al. (2000), Rahman (2000), Liu et al. (2001), Ali et
al. (2002) and Ranasinghe et al. (2002) identified that fungi are influenced by the
essential oils from a number of species in the genus Cinnamomum.

It is not the intention here to discuss the allelopathic possibility of all the biochemical
components within the camphor laurel tree as the technique of gas chromatography
uses volatile oils to differentiate various chemo-types, and does not provide a full
identification of possible allelochemic compounds, as Rice (1984) indicates to
possibly involve many plant biochemicals.

The camphor tree rapidly forms dense stands which dominate large areas of open
land, disturbed areas or regenerating forests (Firth 1979, Schenk and Wallace 1996,
Neilan 2004,) often with a deep layer of slowly decomposing leaf litter (author pers.
obs.). Through the process of rainwater and fog leaching of fallen and living leaves
and branches as well as release from the shallow and extensive root system, a pathway
for possible allelopathic activity below the tree can be envisaged. What may facilitate
allelopathy is the presence in the tree of a dense canopy which shades the soil surface
from bright sunlight and an extensive, shallow and competitive root system which
may influence soil nutrient and moisture availability in the upper horizon, factors
which Whittaker (1970) indicate exacerbate allelopathy. The tree is also observed to
have a high above ground competitive ability with rapid growth of seedlings, high
biomass accumulation in the mature tree (Neilan 2004) and a high longevity, perhaps
thousands of years (Pakenham 2003). What this may also indicate is that the tree has
potential to store large amounts of allelochemicals throughout its structure and exert
any possible allelopathic effect on the surrounding plants for very long periods of
time. Considering also that two litter falls occur per year in the region (author pers.
obs.), resulting from the process of vernal abscission (Addicott 1978) and slow litter
53

decomposition (author pers. obs.), the potential of this tree to exert allelopathy on the
surrounding plants and hence influence plant succession may be real.

An alpha selection may possibly occur if allelopathy is identified as active in the field
situation for this tree. Differences may also result in this selection as a result of
variation in oil composition of the chemo-types. This may produce changes to plant
succession with those species more tolerant of the allelopathic effect or stimulated by
it, becoming more dominant. This also requires the possible effect to be separated
from any effect resulting from the plants competitive ability i.e. possible beta
selection. However, this requires further testing to verify whether allelopathy is
significant in the ecology of the species and if any differences in effect may exist
between the two chemo-types.

1.3.4 Research Hypothesis and Approach

As a result of the complex biochemical release of many terpenoids and phenolics of


camphor laurel through possible rain-wash and fog drip from the canopy, leaching and
decomposition of fallen leaves, twigs and branches, root exudation and rain-wash
from the trunk, it is unknown what phytotoxic effect these varied allelopathic
compounds may have on the way native forests regenerate under a camphor laurel
canopy in the Big Scrub region. The structure of this research was aimed towards
obtaining an understanding of how active allelopathy may be in influencing forest
regeneration in areas dominated by the tree and what role other capability attributes
may have in enabling the tree to exert any possible allelopathy.

The research hypothesis (H0) is:


Camphor laurel exerts no significant effect on the regenerating plant assemblage of
the local region due to its ecology, and

the release of possible allelopathic compounds from the aqueous leaf leachates of the
two chemo-types, 'cineole' or 'camphor' do not contribute to the trees success as an
exotic intrusive element in the flora.

54

The geographical focus of the study is the north-eastern NSW, Big Scrub region due
to extent of the invasion of camphor laurel, and the high diversity of species and
vegetation types present. The location provides for a high camphor laurel to native
vegetation interaction, and a range of species for testing.

The research is structured in the following manner:

Review of Camphor Laurel Ecology (Chapter 1): A literature

review of what is known worldwide about the trees ecological strategies and the
status it has reached as an intrusive exotic plant within Australia;

Allelopathic Seed Germination and Algal Growth Trial (Chapter

2): A glasshouse germination and growth trial involving the application of aqueous
leaf extracts of the two locally known camphor laurel chemo-types, 'cineole' and
'camphor', to a broad range of local vascular plant seed and soil algae to establish
the importance of allelopathy in the germination and growth of the regeneration
assemblage;

Leaf Morphology and Olfactory Recognition of the 'Cineole' and

'Camphor' Chemo-types (Chapter 3): A morphological comparison of the leaves of


camphor laurel as an aid in differentiating the two locally known chemo-types;

Field Assessment of Allelopathy in Vascular Plant Seedlings

(Chapter 4): A field analysis comparing species from the allelopathy glasshouse trial
to the field situation, to establish if there is verification of trends in growth as a result
of any allelopathy in the field; and

Camphor Laurel: Potential and Possibilities (Chapter 5):

Importance of the findings for the long-term viability of regenerating forests in northeastern New South Wales including recommendations for possible control measures,
management and research.

55

1.3.5 Significance of the Research


Results obtained from an allelopathy trial on a variety of native regeneration plants
and exotic weeds exposed to camphor laurel leachates, and a field survey of plant
regeneration under the two locally known chemo-types, will contribute data necessary
for understanding the possible long-term allelopathic impact of this tree on the
successional sequence in vegetation invaded, and the role that allelopathy may play in
the ecology of the tree. Understanding the behaviour of germination and recruitment
in various native seeds and weeds will give an indication of the significance of
camphor laurel allelopathy in manipulating plant succession. It will also identify
whether allelopathy is important in altering or suppressing growth in the plant
assemblage for some species while others may benefit.

Overall, this knowledge will contribute to an understanding of the invasive impact of


camphor laurel on native plant communities i.e. How camphor laurel manages to be
successful as an invader; whether the process of invasion changes successional
processes in invaded communities, and if this is a threatening process to some native
species or assemblages in the camphor laurel invaded regeneration.

In a management context it will contribute to a better understanding of the plants


ecology, particularly the process of invasion, dominance and inhibition which will be
important in establishing its future weed status, including control or cultural methods
deployed during regeneration in camphor laurel communities.

The research also offers an opportunity to provide a direction for further studies and
experiments with camphor laurel effects on the plant community, particularly leachate
influences on soil and water organisms and its impact on water quality.

56

-Chapter 2:The Influence of Aqueous Leaf Extracts of Camphor


Laurel on the Germination of Seed and
the Growth of Soil Algae.

2.1 Introduction
Asplund (1968, 1969) found that the monoterpenes, some of which are commonly
found in camphor laurel, repressed the germination of vegetables and field crops.
Research into the allelopathic influences on seed germination using camphor laurel
extracts was undertaken by Firth (1979), Chou et al. (1989) and Johns (1994). These
studies identified a variable effect on the seed germination of a small number of
exotic trees, grasses, vegetables and native Australian plants. Visual observation by
Scott (1999) suggested that camphor laurel allelopathy had little effect on the
regeneration of plants in north-eastern New South Wales and indicated that light,
moisture and nutrients are responsible for the development of stunted seedlings below
camphor laurel. However, Paul et al. (2010) in a study of seedling growth in soil
below camphor laurel found a 25% reduction in the growth rates of three native
species associated with regeneration in the Big Scrub region.

Detailed effects of camphor laurel extracts on seed germination of native species and
soil algae associated with these communities have not been assessed. Therefore, little
is known regarding the long term impact of aqueous leaf extracts of camphor laurel on
the composition of the regenerating assemblages of the region. Do extracts of
camphor laurel influence native regeneration species, weeds of consequence and algal
composition of the regenerating plant assemblages? Can it ultimately affect soil
processes, plant growth and the food supply for leaf litter and soil organisms?

57

The following experiment is an attempt to establish if germination and growth of


vascular seedlings from a representative sample of the flora in north-eastern NSW,
and soil algae are being influenced by aqueous leaf extracts of camphor laurel under
glasshouse conditions. This will establish a basis for field assessment of any identified
effect, through providing information on the proportion of species possibly
influenced, how this may be occurring, and whether individual species that show
significant response are being stimulated or inhibited.

2.2 Aim and Hypothesis


The aim of this experiment is to establish whether the aqueous leaf extract of camphor
laurel has:

an allelopathic influence on the germination of a variety of common native


plants and woody weeds and soil algae in the camphor laurel regeneration
assemblages, and

whether any significant differences in allelopathic effect on germination and


early growth of these species is effective for the 2 locally known chemo-types i.e. the
cineole chemo-type and the camphor chemo-type.

Therefore, the null hypothesis H0 being tested is:


there is no significant effect on the germination and growth of vascular
plants and soil algae from the regeneration assemblages under glasshouse conditions
resulting from camphor laurel leaf extracts, and

there is no significant difference in allelopathic effect due to the chemo-types


on seed germination, early plant growth and algal populations from the assemblages.

2.3 Methods
2.3.1 Experimental Design
The experimental design included 2 replications of 3 treatments (including a control)
for each of the vascular species and algal samples trialed. The 3 treatments consisted
58

of aqueous extracts of the 2 known local camphor laurel chemo-types i.e. cineole
and camphor and a control consisting of tap water. The control had pH, Total
Nitrogen (TN) and Total Phosphorus (TP) adjusted to mid-point between the two
extracts, being important macro-nutrients utilized by the early growth of plants. The
use of more than 2 replications was avoided due to space limitations and the large
number of species tested. The experiment was not repeated as previous work
undertaken by Firth (1979), Chou et. al. (1989) and Johns (1994) demonstrated
allelopathic effect and repeating the work could not be achieved within the time
constraints of the research. The solutions were collected, prepared and applied
according to the methods identified below.
2.3.2 Experimental Unit and Volatile Loss
In an early seed experiment by Firth (1979), Petri dishes containing extract were used
but with the possible loss of volatiles which may have been the active allelopathic
compounds. This was identified as the possible reason why this earlier experiment did
not produce significant results, and was rectified in a second experiment where closed
lids were used.

In this experiment precautions were taken to minimize volatile loss through the
hermetic sealing of all extract storage drums, the use of pressurized sprayers and the
construction of a specialized propagation unit in the glasshouse. This unit was
designed not only to reduce possible volatile loss but to also avoid cross
contamination of extracts or volatiles, to minimize excessive mist-spray entry and
resultant washout or dilution of extracts and to provide evaporative cooling so as to
closely mimic the moist humid conditions of a leaf litter environment.

The units (Figure 4) were constructed from polystyrene vegetable boxes of 40 x 55 x


30cm and covered with Poly-shade of 85% light entry. However, the boxes were
not entirely covered with the Poly-shade as a 1cm gap at each end was provided to
allow for the ventilation of excess heat, and allow entry of moisture from mist drift
without the washout of extracts occurring from the punnets. Initially, the punnets were
placed onto a 2cm bed of sand in the unit with drainage holes provided along the
sides. This was replaced with plastic pipe following the observation of water-logging
in the first few weeks of the trial, due to silt and algal accumulation in the sand. The
59

propagation units were also grouped separately according to the treatment being
applied, with different treatments being located a metre distant from each other to
avoid possible cross contamination. Replicates were also grouped together but in
spacer pipe to allow
moisture shedding from
Polyshade
vent and 1cm gap
for moisture entry
Polyshade 85%
light entry

55cm

polystyrene
vegetable box

Polyshade
taped to box

clip

40cm
drainage holes
plastic pipe
for aeration
under punnet

punnet

30cm

Figure 4. Propagation unit used for the germination of seed and growth
of algae in the allelopathy glasshouse trial.
separate propagation units. Over time the number of propagation units per treatment
reached 6 at full capacity i.e. 18 units in total, containing 12 punnets each, or 216
punnets in total.

In further work by Firth (1979) acid-washed coarse river sand was used as the
medium and extracts prepared from leaves and roots of the camphor laurel tree. Johns
(1994) also used two parts washed coarse river sand to one part peat moss, Chou et
al. (1989) sandy loam, and Panetta (2000) a commercial topsoil mixture. In these

60

experiments where sand or topsoil mixtures were the selected media for germination,
allelopathy was identified as being active indicating that both sand and commercial
topsoil media are suitable for use in allelopathy trials with camphor laurel extracts.

This experiment used washed river sand for the germination of the vascular plants and
fine white building sand for the growth of algae. Punnets of 8 x 14 x 4.5cm were used
for propagation. For the vascular plants 50 seeds planted per replicate providing 100
seeds per treatment for the control, camphor and cineole applications, providing
statistical validity. Algae were inoculated onto the punnet surface using a culture of
soil algae diluted in tap water.

2.3.3 Environmental Parameters


The propagation units were installed in an unheated glasshouse in an area of light
misting for a 12 month period from early summer 2003.

Light levels varied during the day and month across the propagation area and a
routine of weekly rotation was implemented to ensure that all units received similar
light and mist levels.

Temperature was recorded within the propagation unit as maxima and minima during
the experiment on a regular basis i.e. whenever seedling measurement occurred. This
was commonly each day, but with a maximum outside limit of 3 days between
recordings. During winter the time between recording periods was reduced to 3 days,
as temperatures were reaching minima of 5C, which resulted in slower germination
and evaporation from the punnets.

Although environment control was not available in the glasshouse, temperature was
ameliorated to some extent through the use of misting, shade-cloth and placement of
the units in the coolest or warmest positions depending on seasonal temperature
variability. Initially, the propagation units were placed in the coolest location i.e.
under mist on the glasshouse floor, when temperatures were found to exceed 45C.
This enabled maximum temperatures to remain below 45C until mid-summer when
shade-cloth of 60% density was installed above the units permitting further control
over higher temperatures. This was removed in late autumn when temperatures
61

reduced to maxima of 27C. At this time the propagation units were moved upwards
onto benches where conditions were warmer in preparation for winter. This strategy
allowed some degree of control in both very hot and cooler conditions, providing the
best possible growth for germinating seed under a limited glasshouse environment
control.
2.3.4 Extract Preparation and Application
Firth (1979) used 30 grams of camphor laurel roots and leaves soaked for 3 days in 1
litre of de-ionised water for seed treatments. This extract was applied to media free
Petri dishes 3 days before planting, at planting and 3 days after planting, and a sealed
lid applied. Chou et al. (1989) used air-dried leaves of the camphor laurel tree through
aqueous extraction resulting in a series of concentrations of 1%, 2%, 3%, 4% and 5%,
where 1% is equivalent to 10 grams of dried leaves to 1 litre of water. The extract was
then filtered through Whatman 42 filter paper and stored at 5C. Johns (1994)
prepared camphor laurel extract by taking 200 grams of leaves and preparing a slurry
in a food processor for 5 minutes. This was added to 4 litres of water and allowed to
stand for 4 hours before further dilution to 9 litres giving a final preparation
proportion of approximately 22 grams of material per litre. The extract was applied
every two days. Allelopathic effect was recorded with the application rates and
methods used by Firth (1979), Chou et al. (1989) and Johns (1994).

A pilot trial to determine the most effective rate was not determined as Chou et al.
(1989) found that concentrations ranging from 1% to 5% all produced effect. The
proportion of 30 grams of camphor laurel leaves per litre was adopted for this
experiment, enabling comparison against previous experiments identified above. This
was achieved through the collection of 900 grams of mature camphor laurel leaves,
450 grams from each of 2 known trees of the cineole as well as the camphor
chemo-types at Casino. These were identified previously by Stubbs (pers. com. 2003)
through gas chromatographic analysis (Stubbs and Brushett 2001). Leaf material was
processed to a fine cut in a food processor with tap water, and further diluted with 20
litres of water for the 2 chemo-types respectively, then soaked for 3 days. Following
soaking, the extracts were filtered into hermetically sealable 30 litre plastic drums
using a 63m nylon gauze following mucilage blockage of No.1 to No. 8 filter papers.
After filtering into the storage drums, a further 10 litres of tap water was added to
62

bring the volume to 30 litres. Due to the many phenolic and terpenopoid compounds
indentified in Chapter 1 as possible bioactive constituents in camphor laurel, the
varying ways these behave on exposure to the environment and the differences in
chemical composition between the chemo-types it would not have been practical to
select one compound to determine the relevance of the work in the field situation.
This work therefore, reflects the synergistic effect of the many compounds in each
individual chemo-type.

During the process of extract preparation a control solution tank was also filled with
water which provided the 3 vats with water of the same age, in an attempt to alleviate
possible variability in volatile loss of chlorine derived from the tap water. The 3
solutions of cineole, camphor and the control were then analyzed for pH, TN and
TP. The pH of the solutions was assessed using a calibrated pH with pH 4 and buffers
7, prior to each use. TN and TP were analyzed in the Southern Cross University
(SCU) Environmental Analysis Laboratory (EAL) following submission of 3, 10mL
undiluted samples of the solutions prior to the adjustment of the control.

The control was then adjusted for pH, TN and TP following molarity calculation for
TN and TP to fall between the concentrations of the leachate solutions. TN and TP
were adjusted respectively with mono-ammonium phosphate (MAP) NH4H2PO4 and
urea (NH2)2CO following the molarity calculations. A 2 molar solution of HCl was
used to gradually adjust the pH using an auto-pipette following TN and TP
adjustment.

All control solutions were submitted to the EAL prior to and following chemical and
pH adjustment to establish consistency of pH, TN and TP between the prepared vats.
Any errors were recalculated, re-adjusted and resubmitted to the EAL until
consistency was attained.

Pressurized sprayers were used for the application of the three solutions three times
per week on average. The pressurized containers provided for the continual sealing of
the solutions from the atmosphere, reducing possible volatile loss and oxidation, in
addition to allowing even and controlled application of the treatments over the planted
punnets, with minimal disturbance to the media. The vat solutions were stored in the
63

laboratory at a temperature of 21-25C for the period of the experiment and were
tightly sealed.
2.3.5 Vascular Plant Selection, Treatment and Planting
The criteria for the selection of local species for the experiment should ideally be
based on a sampling of a diverse subset of the common plants associated with various
plant assemblages invaded by camphor laurel in the region i.e. regenerating plant
communities. Survey work in 179 remnants in the region, 32 of which were
dominated by *C. camphora, identified a large number of tree and shrub species
associated with camphor laurel communities (Schenk and Wallace 1996). These
species formed the initial selection of the target groups of interest for testing. To add
further life-form groups to these initial target groups, species other than trees and
shrubs were also selected being based on field observations in the camphor laurel
assemblages. These supplementary target groups not represented in the initial plant
survey included vines and herbaceous plants of the lower stratum.

Most seeds needed some preparation prior to germination. For example, all fleshy
fruited species required pericarp removal, and species that had hard seed coats such as
Acacia melanoxylon, and Commersonia bartramia required hot water scarification
and overnight soaking to break physical dormancy. Fine seeds were handled with a
small aspirator following counting and sorting under a low powered microscope to
divide chaff and potentially viable seed. Small seeds were then placed onto a grid
system 2mm below the media surface of the punnet using a moistened probe. Larger
seeds were de-pulped and washed with tap water prior to being placed at 1cm below
the media surface on a grid system. Cuttings were also prepared for a few herbaceous
species such as Pratia purpurascens and Viola hederacea, and for *Anredera
cordifolia, aerial tubers were utilized. In the case of P. purpurascens the plants were
cut into 2cm lengths with all leaves removed, and planted vertically 1cm deep into a
grid within the punnet. For V. hederacea all leaves and auxiliary roots were removed
to the bud and tap root before being planted into the punnet grid. With *A. cordifolia
however, the aerial tubers were subdivided into approximately 0.5g portions
containing a large dormant bud and no leaves.

64

Emergence was recorded for the vascular plants when the hypocotyl appeared at the
soil surface, while for cuttings initiation of growth was established when auxiliary
buds began growing. Germinating seeds were tagged to avoid recounting and an
accumulated total recorded at each visit. A tetrazolium test was not attempted for
seeds that did not germinate as many seeds were either microscopic, very small or
difficult to locate within the media.

All plants from which seeds were collected are vouchered through the Southern Cross
University herbarium. Specimens were fully pressed, mounted and labeled. In the
identification of the vascular plants to species level the following plant keys and floras
were used: Williams, Harden and McDonald (1984), Harden (1992), and Bale (1992).
Scientific names adopted in the study are according to CSIRO (1999).

2.3.6 Algae Selection, Inoculation and Treatment


The inclusion of soil algae in the experiment was initially co-incidental. During the
first month of the trial it was noted that soil algae growing in the media of the seed
germination trial were more abundant and differed in appearance in the control than
the camphor and cineole treatments. It was decided to test the soil algae based on
this visual observation.

Samples of 5g of algae from the glasshouse floor which was identified to have high
diversity, and two cryptogamic mats from wet sclerophyll forest near Uki, NSW were
used for inoculation in 3 separate algal trials. Algal scrapings from the glasshouse
floor were used as this was identified as possessing high diversity following
microscopic examination. The sample was removed from the original substrate,
placed in 1 L of tap water and shaken vigorously until dispersion was obtained in the
solution. This was then filtered through coarse nylon gauze and applied over the
punnets using a hand sprayer. Identifications of the individual soil algae were
performed on the first algae trial (sample 1) only, as identifications and processing of
the material was time consuming due to the incomplete nature of the taxonomy of the
Australian soil algae. The identifications were based on Prescott (1978), Bold et al.
(1980), Scagel et al. (1980), Entwisle et al. (1997) and Baker and Fabbio (2002) with
identifications to generic level. Where this was not possible due to algal genera not
65

being included in the key, undescribed species, or species difficult to identify, they
were classified to Division and labeled with a species number.

2.3.7 Data Types Collected


For the vascular plants, data collection involved 3 forms: pre-emergent measurement
during seed preparation, emergence measurement during germination, and post
emergence measurement following the completion of germination to identify any
differences in vascular plant growth.

Specifically the data collected included:


Pre-emergence Measurement:
The seed was initially identified as either having a moist or dry pericarp. Once the
pericarp was removed from the seed and dried with a paper towel, the weight of one
hundred seeds was obtained for each species trialed.
Emergence Measurement:
Species, treatment type, date planted, daily count of accumulated germination,
germination completion date, and notes on, for example, discolouration, disease,
death, dormancy, were collected throughout the germination of the species i.e. until
germination plateaued in all treatments.
Post Emergence Measurement:
Species, treatment type, date, measurements of shoot length, leaf number (excluding
cotyledons), radicle or root length, and notes on seedling health, cotyledon health,
discolouration or necrosis, disease and seedling deaths were gathered following
removal of the seedlings from the punnets.

For the soil algae where area covered was measured during each recording period, a
grid was constructed of 400 1cm squares on plastic sheets. These were overlaid onto
the punnets for recording and sequentially marked off as the cryptogamic mat
developed over the media. Following full growth coverage of the punnets by the
66

algae, 6 samples of 3mm2 were taken randomly from each of the 3 treatments and
mounted on slides. These were examined under low and high power magnification
(40x to 1000x), with genera being identified and grouped on an abundance scale of 1
to 3; with 1 being absent, 2 being present but not common, and 3 being common to
abundant. An abundance score was calculated through the addition of the scores for
each species and division by the mean number of samples for each treatment.

2.3.8 Statistical Approach


Results for the germination and algal growth trial were initially graphed to provide a
visual comparison of emergence and treatment effect over time for each species. A
non-linear regression using the Richards function (Richards 1959, Bertalanffy 1960,
Cheng and Gordon 2000) was performed on each of the three curves represented for
each species treatment using Excel Solver, as the Richards model best represented
the sigmoid growth displayed by the graphs. The modeled output from the function
provided an upper asymptote, mid-inflection point, days to mid-inflection and
growth rate for each treatment using the Excel equivalent of the Richards (1959)
formulae:
L = A(1e(-kt))-1/v ,

where L is the germination or algal coverage percentage at t(d), A is the upper


germination or algal coverage asymptote, v estimates the curve inflection point, is
the x-axis placement parameter (time) and k is the rate of change parameter.

The Excel equivalent to the Richards formulae was:

Y = B2/(Power((1+(C2*Exp(-D2*(A5-E2)))),1/C2))

where Y is the modeled data point, B2 is the asymptote, C2 is the location on the xaxis (time in days), D2 is the scaling factor (germination rate), A5 is the position on xaxis of the observed data (days), E2 is the time (days) to maximum growth inflection
point and 1/C2 is the calculated position on the y-axis as percentage germination or
coverage. Standard errors were not possible to generate in Excel Solver but would
67

have been possible using SPSS. However, SPSS suffered from an inability to
accept the Richards function and could not be applied.

These modeled data were then statistically compared in a deviance test (loglikelihood
ratio) and multi-level analysis using the software MLwiN. The asymptote, midinflection point and germination rate were approximately randomly distributed. For
the days to the mid-inflection point parameter distribution was right-skewed and the
data transformed to natural logarithms which randomized the distribution.

Multi-level models (Snijders and Bosker 1999), with all plant species and treatments
being the control, cineole and camphor extracts within species as nested random
factors and 2 dummy variables for the 3 treatments as a fixed factor were fitted to the
data. Repeated mean likelihood (REPIL) estimates providing a maximum likelihood
fit were obtained using MLwiN. The significance of the treatment effect was
assessed by the likelihood ratio test where the difference between loglikelihood
statistics of the intercept only model and the model including the treatment factor is
tested against a chi square with 2 degrees of freedom (Snijders and Bosker 1999).
Residual plots were calculated for each of the 4 parameters of asymptote, natural log
of the days to mid-inflection, mid-inflection point, and germination rate or
coverage rate with an sd of +/- 1.4 as outlined by Goldstein (1995).

The post-emergent data for the vascular plants collected at the completion of the trial
included shoot length, radicle length, leaf number and leaf area of the largest leaflet of
all seedlings. The data were statistically compared using a two-sample t-test assuming
equal variance between the control and the cineole treatment, the control and the
camphor treatment and the cineole and the camphor treatment. This provided for
comparison of effect between the control and the other two treatments as well as any
differences between treatments.

Excel was used to perform the statistical calculations for vascular plant germination
and algal growth over time through the application of macros written to accommodate
the Richards function (Richards 1959, Bertalanffy 1960, Cheng and Gordon 2000).
Using the programming techniques outlined by Liengme (2000), Excel Solver was
applied to fit the Richards curve using precision set at 0.000001, tolerance at 5%, and
68

convergence at 0.0001 with a Newton algorithm. Starting points for zero values were
set to 0.0001 as a zero value is not accepted by the function. This provided not only
fitted germination, SS differences and residuals for the dataset but most importantly r2
values and four modeled data points, i.e. upper asymptote, mid-inflection point,
days to inflection and growth rate, for input to MLwiN. The use of the statistical
software SPSS was also attempted but suffered from the inability to allow the use of
the Richards formulae without lengthy experimentation. Data sets in the case of SPSS
would have needed to be broken into component sections to allow for statistical
analysis, thereby losing some continuity due to a disjunct analysis and creating further
statistical calculations on an already large data set.

Knusel (1998), McCullough (1998, 1999), McCullough and Wilson (1999) and Cox
(2000) have expressed concern over the accuracy of Excel for statistical application.
The assessments by Cox (2000) considered the calculation of distributions (tail
probabilities), mean and standard deviation, analysis of variance, linear regression,
non-linear regression (using Excels Solver) and random numbers. Many of these had
also been previously tested by Knusel (1998), McCullough (1998, 1999) and
McCullough and Wilson (1999). Coxs work found that Excel did not perform as well
as SAS, SPSS or S-Plus especially with non-linear regression due to nave
algorithms, rounding and truncation errors but in extreme situations only, where
numbers exceed fifteen digits for Excel versions later than the year 2000. In the
analysis of distributions, rare cases resulting in failure do occur where extreme tails
are beyond 10-6, but again this does not often occur in significance testing. Cox (2000)
suggests that if failure does not occur, the result can be regarded as being reliable for
distributions. For the analysis of variance Excels nave algorithms and rounding
errors can create problems where very large numbers are present. However, when
compared with SAS (Anova) and SPSS these programs did no better with large
numbers. Excels Solver was found to perform well providing results similar to other
statistical packages. The problem with Solver lies in its inability to provide standard
errors with the estimates and so requires verification. Overall, Cox (2000) found that
when using Excel for simple summaries and tests such as t-tests, chi-square and
regression analysis Excel will give the correct answer. Now that Excel uses fifteen
significant digits instead of seven the impact of poorer algorithms is much less.

69

In this study of phytochemistry, analyses in Excel that were performed did not
transcend the inadequacies identified for the program i.e. data or statistical numbers
did not exceed fifteen digits, and the output from the regression using the Richards
function was checked against separate calculations of the residual values and
manually against the original dataset for inconsistencies.
2.3.9 Vermin Control and Fungal Invasion
Slugs and cockroaches were observed to browse young seedling shoots, rats predated
seed and ants removed seeds with attached elaiosomes or the scant remains of sweet
pericarp. Vermin control during the experiment included the use of various chemical
baits to control these pests. The chemicals included Defender (metaldehyde) for
slugs, Racumin (coumatetralyl) for rats, Nest Kill (chlorpyrifos) for cockroaches
and Antex (chlorpyrifos) for ants.

For the control of ants, slugs and cockroaches, baits were laid within the propagation
units themselves and placed on the floor of the unit to prevent contamination of the
germination punnets. This was repeated every month during periods of predation. Rat
baits however, were strategically placed throughout the glasshouse at floor level and
re-application periods were based on observation of disturbance to the baits. This
replacement was usually fortnightly during periods of rat predation.

During early to mid summer, fungal invasion of the seedlings was noted with the
initial growths occurring in the cineole and camphor treatments, but latter
spreading to the control. This was identified as Rhizopus spp., a Zygomycete, in its
sexual stage. The fungus was found to be of saprophytic nature and did not result in
any infestation or death of any seedlings. The species was also observed in its asexual
stage floating on the top of both the cineole and camphor extract tanks. This
indicates that it may have originated from the phylloplane or tissues of the camphor
laurel leaves used for the preparation of the solutions. The growth however, did
eventually subside as temperatures in the glasshouse began decreasing in early
autumn.

70

2.4 Results

2.4.1 Propagation Unit Temperature


Internal propagation unit temperature collection (Figure 5) occurred for the full period
of the experiment i.e. 355 days starting in spring 2003 and ending in spring 2004.

Maximum temperatures during the spring-summer period ranged between peaks of


47C to 19C under cloudy conditions. The autumn-winter maxima however, did not
fall below 23C. Minimum temperatures recorded varied between 29C in midsummer to 5C in mid winter.

50

propagation units to floor

shade installed

propagation units to benches


shade removed

45

Maximum

Temperature (C)

40
35
30
25
20

Minimum

15
10
5
0
1

13

28

spring
12 October 2003

46

60

74

89

105

120

135

149

161

summer

173

187

201

215

235

autumn

Day / Season

255

279

319

349

winter

7 October 2004

Figure 5. Propagation unit temperature on a daily and seasonal basis.

2.4.2 Chemistry of the Vat Solutions


In total, 4 vat solution batches were prepared during the trial. Analysis of Total
Nitrogen (TN) and Total Phosphorus (TP) in the control cineole and camphor
solutions for the first 3 vat batches undertaken by EAL identified levels in mg/L. The
fourth batch of solutions used the averages between the second and third batches for
adjustment of TN and TP in the control.

71

The TN and TP levels and completed chemical adjustments for the control solution
in the 3 vat batches are shown in Figure 6a and b; demonstrating the adjusted
consistency between the extract solutions and the control during the experiment.

The pH levels of the 4 batch solutions prior to and following adjustment are shown in
Figure 7a to d.

(a).
14

13.11

12

10.52

10

8.48

mg/L

10.95

9.99

9.49
8.68

8.87

Vat 2

7.38

Vat 1
Vat 3

6
4
0.93

2
0.85

0.69

0
Cont rol- prior t o
adjust ment

Cont rol-adjusted

Camphor

Cineole

solution prepared

(b).
9 .75

10

8 .73
7.8 3

8 .12

7.3 6

Vat 1

mg/L

6 .4 8

Vat 2

Vat 3

4 .72
4 .2 7

4 .0 8

2
0

0 .0 5 0 .0 3 0 .0 1
C o nt ro l- p rio r t o
ad just ment

C o nt ro l- ad just ed

C amp ho r

Cineo le

s olution pr e pare d

Figure 6. Total nitrogen (a); total phosphorus (b) of the vat solutions prepared
for the control, cineole and camphor treatments applied to seeds.

72

(a).
9

pH adustment Control

pH adjustment Control

Control
Camphor
Cineole

pH

6
5
4

solution not in use

solution in use

2
1

10 12 17 21 23 25 28 31 35 37 39 41 44 49 51 55 58
Day

(b).
9

pH adjustment Control

pH

Control

Camphor

Cineole

solution not in use

solution in use

2
1

11 15 20 22 26 29 35 38 44 50 54 56 58 63 69 73 76 83
Day

(c).
9
8
pH adjustment Control

pH

pH adjustment Control

Control

Camphor
Cineole

solution
not in use

solution in use

2
1

8 12 14 18 21 22 27 28 33 34 40 41 53 56 60 62 63 64 66 67 69 70 73 78
Day

73

(d).
9
pH adustment Control
8
pH adjustment Control
7

Control

pH

Camphor
Cineole

5
4

solution in use day two

3
2
1

12

15

22

26

28 30
Day

34

39

41

43

45

49

55

60

63

Figure 7. pH adjustments of the four vat solutions prepared for the treatments
applied to seeds for: (a) Vat solution 1; (b) Vat solution 2; (c) Vat solution 3;
(d) Vat solution 4.

2.4.3 Vascular Species Trialed


The vascular species trial assessed 54 species associated with camphor laurel
regeneration assemblages in north-eastern NSW, which included 31 native trees and
shrubs, 6 introduced trees, 2 native vines, 3 introduced vines, 8 native herbs and 4
introduced herbs.

A list of trialed vascular species and their seed characteristics are shown in Table 6,
together with Family, life-form, pericarp moisture, observed dormancy and seed
weight.

The vascular species target groups used for seed collection were also compared
against the species actually trialed, with 44 of the 54 trialed plants i.e. 81%, being
represented as a target species. Further comparison to plants observed by Neilan
(2004) in the camphor laurel assemblages also revealed that 33 or 61% of the trialed
species also occurred in that survey.

2.4.4 Germination Pre-statistical Results


The pre-statistical analysis of vascular plant germination response to the extracts are
discussed in 2 sections i.e. germination response of the 2 extracts compared with the
control for individual species; and responses to the individual extracts for species,
family, life-form, seed weight, seed moisture, introduced species, and dormancy of
seed.
74

Table 6. Vascular plants trialed and their seed characteristics arranged by seed weight.
Species

Family

Life-form

Pericarp Moisture

Observed
Dormancy

100 Seeds
Weight (g)

dry
dry
dry
dry
dry
dry
dry
dry
dry
dry
dry
dry
dry
moist
dry
dry
moist
dry
dry
moist
dry
dry
dry
dry
dry
moist
moist
moist
moist
moist
dry
moist - veg. propagation
moist - veg. propagation
moist
dry
moist
moist
moist
moist
moist
dry
dry
moist
dry
moist
moist
moist
moist
dry
moist
moist - veg. propagation
moist
dry

N
N
N
N
N
N
N
N
N
N
N
N
N
N
D
N
N
N
N
D
D
N
D
D
N
D
D
D
D
D
N
N
N
N
D
N
D
N
D
N
N
N
N
N
N
D
N
N
N
N
N
N
N

0.008
0.01
0.01
0.013
0.02
0.02
0.07
0.07
0.07
0.08
0.08
0.09
0.09
0.15
0.25
0.31
0.33
0.39
0.4
0.42
0.5
0.55
0.8
1.5
1.79
1.85
1.87
2.12
2.31
2.77
3.25
3.32
3.57
3.74
4.5
4.6
5
5
5.24
7.38
7.58
7.79
11
12.08
12.77
14.4
19
22.14
30.33
40.07
52.52
77
4790

Callistemon viminalis

Myrtaceae

*Cyperus enervis

Cyperaceae

Oplismenus aemulus

Poaceae

Ottochloa gracillima

Poaceae

Lophostemon confertus

Myrtaceae

Eucalyptus saligna

Myrtaceae

Allocasuarina torulosa

Casuarinaceae

Eucalyptus microcorys

Myrtaceae

*Setaria sphacelata var. narok

Poaceae

Helichrysum bracteatum

Asteraceae

*Paspalum dilatatum

Poaceae

*Chloris gayana

Poaceae

*Pennisetum clandestinum

Poaceae

Ficus coronata

Moraceae

Commersonia bartramia

Sterculiaceae

Allocasuarina littoralis

Casuarinaceae

Ficus macrophylla

Moraceae

Tristaniopsis laurina

Myrtaceae

Toona australis

Meliaceae

Cordyline rubra

Agavaceae

Hymenosporum flavum

Pittosporaceae

Eucalptus intermedia

Myrtaceae

Acacia melanoxylon

Mimosaceae

Lomandra longifolia

Xanthorrhoeaceae

*Solanum capsicoides

Solanaceae

*Lantana camara

Verbenaceae

Geitonoplesium cymosum

Philesiaceae

th

*Passiflora suberosa

Psssifloraceae

th

*Ligustrum lucidum

Oleaceae

Alpinia caerulea

Zingiberaceae

*Cassia coluteoides

Caesalpinaceae

Pratia purpurascens

Lobeliaceae

Viola hederacea

Violaceae

Omalanthus populifolius

Euphorbiaceae

Mallotus philippensis

Euphorbiaceae

Macaranga tanarius

Euphorbiaceae

Alphitonia excelsa

Rhamnaceae

Guioa semiglauca

Sapindaceae

Cissus hypoglauca

Vitaceae

Th

Jagera pseudorhus

Sapindaceae

Flindersia xanthoxyla

Rutaceae

Lepiderema pulchella

Sapindaceae

Syzygium paniculatum

Myrtaceae

*Cardiospermum grandiflorum

Sapindaceae

Th

Syzygium luehmannii

Myrtaceae

*Cinnamomum camphora

Lauraceae

Cupaniopsis parvifolia

Sapindaceae

Diploglottis australis

Sapindaceae

Araucaria cunninghamii

Araucariaceae

Melia azedarach

Meliaceae

*Anredera cordifolia

Basellaceae

Th

Davidsonia pruriens

Davidisoniaceae

Castanospermun australe

Papilionaceae

* - introduced species.
1. Life-form: T tree, S shrub, Th thick-stemmed vine, th thin-stemmed vine,
g graminoid, f forb.
2. Observed Dormancy: N no dormancy, D dormancy (extended period without germination).

75

Throughout the 3 life-form groups (trees and shrubs, herbs and vines) germination of
seed or strike response of cuttings, stolons and tubers was found to fall into 9 groups,
7 of which can be regarded as graphically defined responses to the two extract
treatments compared with the control. The remaining 2 groups were identified as
error or with germination below 10% making comparisons and analysis difficult.

The graphical responses of the cineole and camphor treatments when compared
with the control included:

Group 1: treatments with higher germination than the control;

Group 2: one treatment with higher germination than the control, the other
lower;

Group 3: one treatment with higher germination than the control, the other
with similar levels to the control;

Group 4: one treatment with lower germination than the control, the other
with similar levels to the control;

Group 5: treatments with lower germination than the control; Group 6:


treatments with similar levels to the control;

Group 7: treatments fluctuating in effect compared with the control i.e. a


germination delay followed by an increase above the control;

Group 8: germination below 10%; and

Group 9: error in control or treatments.

The graphs of the vascular species within each of the 3 life-form groups trialed are
shown in Appendix 1: Germination of the Trees and Shrubs (37 species); Appendix
2: Germination of the Vines and Climbers (5 species), and Appendix 3: Germination
of the Herbs (12 species). These have been arranged in each group in descending
order from the highest attained germination rate to the lowest, providing a measure of
the efficiency of germination for each species.

The species treatment response groups together with the highest germination
percentage in the 3 treatments for each species are shown in Table 7 for the trees and
shrubs and Table 8 for the vines/climbers and herbs.
76

Table 7. Percentage germination of the trees and shrubs in the various treatment
response groups.
% Germination in Treatment Response Groups1
Species
E. intermedia
L. pulchella
E. saligna
S. paniculatum
T. australis
G. semiglauca
M. tanarius
M. philippensis
F. macrophylla
C. viminalis
L. confertus
S. luehmannii
*L. lucidum
*S. capsicoides
O. populifolius
H. flavum
*C. camphora
A. torulosa
C. parvifolia
A. littoralis
*C. coluteoides
A. cunninghamii
T. laurina
F. coronata
C. bartramia
A. melanoxylon
E. microcorys
*L. camara
C. rubra
A. excelsa
D. pruriens
M. azedarach
*C. camphora
F. xanthoxyla
C. australe
J. pseudorhus
D. australis

Group
1

Group
2

Group
3

Group
4

Group
5

Group
6

Group
7

Group
8

Group
9

95
95
92
89
88
87
82
19
13
72
36
92
89
89
85
73
28
19
96
46
38
33
20
98
83
72
34
6
1
0
1032
95
89
90
86
49
29

Treatment response groups:


Group 1: Treatments with higher germination than the control;
Group 2: One treatment with higher germination than the control, the other lower;
Group 3: One treatment with higher germination than the control, the other with similar
levels to control;
Group 4: One treatment with lower germination than the control, the other with similar
levels to the control;
Group 5: Treatments with lower germination than the control;
Group 6: Treatments with similar levels to the control;
Group 7: Treatments fluctuating in effect compared to the control;
Group 8: Germination below 10%;
Group 9: Error in control or treatment.
2
Germination of 103% is the result of polyembryony.

77

Table 8. Percentage germination of the vines and herbs in the various treatment
response groups.
% Germination in Treatment Response Groups1
Species

Group
1

Group
2

Group
3

Group
4

Group
5

Group
6

Group
7

Group
8

Group
9

Vines/Climbers
*C. grandiflorum
*A. cordifolia
C. hypoglauca
G. cymosum
*P. suberosa

76
75
83
7
3

Herbs
O. aemulus
C. enervis
*S. sphacelata
O. gracillima
A. caerulea
*C. gayana
V. hederacea
*P. clandestinum
P. pupurascens
*P. dilatatum
H. bracteatum
L. longifolia
1

97
30
22
17
53
42
98
96
92
56
15
83

Treatment response groups:


Group 1: Treatments with higher germination than the control;
Group 2: One treatment with higher germination than the control, the other lower;
Group 3: One treatment with higher germination than the control, the other with similar
levels to control;
Group 4: One treatment with lower germination than the control, the other with similar
levels to the control;
Group 5: Treatments with lower germination than the control;
Group 6: Treatments with similar levels to the control;
Group 7: Treatments fluctuating in effect compared to the control;
Group 8: Germination below 10%;
Group 9: Error in control or treatment.

The rates of the highest overall germination in the 3 treatments are summarized and
compared within each life-form group in Figure 8.

The individual species responses of the treated seed to both cineole and camphor
extracts compared with the control are now discussed in more detail and are grouped
according to their life-form.

78

120
n = 37

% germination

100

n =5

n = 12

80
60
40
20
0
Trees and shrub s

V ines and t winners

Herb s

Lif e-f orm

Figure . X. Comparison of the highest overall germination


Figure 8. Comparison of the highest overall germination
response in the treatments for each species.

response in

the treatments for each species.

In the trees and shrubs, E. intermedia (Appendix 1, Figure A1.1) germination


commenced on day 6 for all treatments with the control at 21%, cineole at 7% and
camphor at 25%. Lower germination was recorded for cineole until day 23 when
germination returned to near the control level. For the camphor treatment,
germination was at a similar level to the control until day 12 when it was then
exceeded. Germination was completed for the control on day 23 at 95%, cineole
on day 14 at 91% and camphor on day 25 at 95% germination.

With L. pulchella (Appendix 1, Figure A1.2) camphor began germination on day 7


at 1%, and cineole and the control on day 10, both at 41%. The cineole treatment
had lower germination than the control during the entire period. In the case of the
camphor treatment, a similar germination rate to the control occurred until day 10,
when germination exceeded the control for the remainder of the period. Germination
was completed for the control and cineole on day 17 both at 88% and camphor on
day 18 at 95%.

For E. saligna (Appendix 1, Figure A1.3) germination started on day 8 for all
treatments with the control at 49%, cineole at 46% and camphor at 40%. At day
12, both the cineole and camphor deviated from the control. This was slightly
below for the cineole and more substantially above for the camphor which were
both maintained for the remainder of the germination period. For the control,

79

germination was completed on day 27 at 79%, for the cineole on day 18 at 71% and
the camphor on day 32 at 92%.

In the species S. paniculatum (Appendix 1, Figure A1.4) germination of all treatments


commenced at day 10, with the control and cineole at 7% and the camphor at 6%.
The camphor remained slightly below the control level until day 14 when it
exceeded that treatment slightly then returned below the control germination again
at day 15. The cineole treatment deviated above the control at day 11 until day 26
when the control germination was completed at 87%. For both the cineole and
camphor treatments, germination plateaued on day 29 at 89% and 80% respectively.

In the case of T. australis (Appendix 1, Figure A1.5) germination commenced on day


7 for the cineole and camphor with both at 1%, while the control did not start
until day 9 at 3%. Both the cineole and camphor germination remained slightly
above the control until day 13 when germination for all 3 treatments was 55%. After
this time however, both the cineole and camphor remained at a lower germination
rate than the control. Germination was completed for the control on day 22 at 88%,
cineole on day 23 at 84% and camphor on day 23 at 77%.

With G. semiglauca (Appendix 1, Figure A1.6) germination commenced at day 5 for


both the control and the camphor treatments at 1% and 6% respectively, while the
cineole commenced on day 6 at 10%. Germination rates were slightly higher for the
camphor which was maintained throughout the germination period. With the
cineole, germination was slightly less until day 10 when similar levels to the
control were attained until germination was complete. Germination was fully
complete on day 17 for all treatments being 76% for the control, 77% for cineole
and 87% for camphor.

Germination of M. tanarius (Appendix 1, Figure A1.7) commenced on day 11 for


both the cineole and camphor at 2% and 4% respectively, while the control
germinated on day 12 at 1%. With the cineole however, germination steadily
increased above the level of the control for the remainder of the germination period.
For the camphor, germination increased above the control slightly, returned to the
control level then fluctuated slightly above and below the control, eventually
80

attaining a similar germination rate to the control. Germination was complete on day
35 at 61% for the control, day 37 at 82% for the cineole and day 38 for camphor
at 65%.

M. philippensis (Appendix 1, Figure A1.8) commenced germination on day 26 for


cineole at 1%, day 32 for the control at 1% and day 45 for the camphor at 1%.
Although the camphor possessed a slightly higher germination mid-point in the trial,
the control slowly increased germination to a similar level at day 97. The cineole
however, increased above both the other treatments at day 37 to 82 when germination
was at 19%. The germination was completed for the control on day 97 at 5% and for
the camphor on day 112 at 7%. The pattern of germination of this species is also
distinctive with a stepped sequence of germination events numbering 4 episodes in
each treatment.

With F. macrophylla (Appendix 1, Figure A1.9) germination commenced on day 10


for both the control and camphor at 1% while the cineole was delayed until day
13 at 2%. The germination behaviour of both the control and camphor were similar
and attained the same germination levels. For the cineole, germination increased
above the other treatments at day 17 and maintained a higher level until germination
was complete. The germination plateaued on day 16 at 7% for the control, on day 32
at 13% for cineole and day 21 at 7% for camphor.

C. viminalis germination was rapid, (Appendix 1, Figure A1.10) starting on day 4 for
both the control and camphor at 4% and 6% respectively, while for the cineole
this was day 5 at 6%. Germination of both the control and camphor were similar
throughout the germination period and attained higher levels than the cineole.
Germination was completed for the control on day 11 at 71%, day 23 at 49% for
cineole and day 16 at 72% for camphor.

In the species L. confertus (Appendix 1, Figure A1.11) germination commenced at


day 8 for all treatments with the control at 4%, cineole at 6% and camphor at 1%.
The camphor attained a lower germination throughout the trial. The cineole
however, fluctuated slightly above and below the control during the germination
period. Completion of germination occurred on day 40 at 33% for the control, day
81

56 at 36% for cineole and day 69 at 22% for camphor. The germination sequence
for this species was also stepped with as many as 8 germination episodes displayed in
all treatments.

For S. luehmannii (Appendix 1, Figure A1.12) germination started on day 10 for the
control at 4% and day 11 for the other 2 treatments at 14% for cineole and 17% for
camphor. Germination remained lower than the control in cineole and camphor
until the final stages of germination when both these treatments returned to near
control levels. Completion of germination occurred on day 18 for the control at
92%, day 29 for cineole at 90% and day 28 for the camphor at 92% germination.

The germination of *L. lucidum (Appendix 1, Figure A1.13) began on day 29 for the
control at 2%, day 33 for cineole at 2% and day 29 at 8% for camphor.
Germination rates were lower than the control for both cineole and camphor with
the camphor being the lower treatment. However, the cineole did eventually attain
the same level as the control. Germination was completed for all treatments on day
55 at 88% germination for the control, 46% germination for cineole and 89% for
camphor. The graph of this species also indicates that germination may not have
been fully completed on day 55 as plateauing of effect had not fully occurred.

With *S. capsicoides (Appendix 1, Figure A1.14) the germination commenced on day
12 for both the control and cineole at 6% and 1% respectively, with the camphor
being delayed until day 14 at 5%. Germination was lower than the control for both
treatments throughout the germination period. However, the camphor did fluctuate
firstly above the cineole until day 25, when it fell below the level of that treatment.
Germination was completed first by the control on day 31 at 89% germination, then
by cineole on day 34 at 73% and finally by camphor on day 36 at 66%.

In the species O. populifolius (Appendix 1, Figure A1.15) germination commenced at


day 14 in the control at 13% and the camphor at 12%, while the cineole
commenced earlier on day 10 at 1%. Germination rates were lower in the cineole
and camphor for most of the germination period with the levels slowly approaching
the control in the final stages of the germination. Completion of germination was

82

attained for the control on day 28 at 85%, for cineole on day 29 at 80% and
camphor on day 29 at 83% germination.

In the case of H. flavum (Appendix 1, Figure A1.16) germination was slow,


commencing on day 45 for both the control and cineole both at 1%, while the
camphor was delayed until day 50 at 1%. Germination throughout the trial was
lower for both the cineole and camphor with the camphor however, returning
slowly to the same level as the control at day 143. The germination was completed
for the control on day 105 at 72%, day 147 for cineole at 36% and day 145 at 73%
for camphor. The sequence of germination is also markedly stepped in this species,
being most pronounced in the cineole and camphor treatments where as many as
12 episodes of germination plateauing occurred. A lengthy dormancy period of 45
days was also noted for this species.

With the seed of *C. camphora (Appendix 1, Figure A1.17) which was removed from
the soil below a camphor laurel tree at Uki, germination commenced in the control
at day 15 at 1% while the cineole started at day 12 at 1% and the camphor at day
11 at 1%. Overall germination was lower than the control for both extract treatments
with camphor being the lowest throughout the germination period. Germination was
completed for the control on day 34 at 28%, cineole on day 53 at 16% and
camphor on day 83 at 7%. This species also displayed a stepped germination in the
extract treatments of up to 7 episodes for the cineole and 4 episodes for the
camphor.

Germination of A. torulosa (Appendix 1, Figure A1.18) commenced on day 8 for the


camphor at 1%, and day 9 for the cineole and the control both at 3%. Overall,
germination remained lower than the control during the entire period of germination
for both extracts. However, at day 20 cineole became higher than the camphor
germination and was maintained for the remainder of the germination. Completion of
germination occurred on day 26 at 19% for the control, day 33 at 11% for cineole
and day 18 at 6% for camphor. Stepped germination sequences also occurred in each
of the treatments of up to 4 episodes, and were particularly pronounced for the
control and cineole.

83

With C. parvifolia (Appendix 1, Figure A1.19) germination started at day 8 for both
the cineole and camphor at 3% and 1% respectively and day 10 for the control at
33%. Similar germination occurred for all treatments with slight deviations above and
below the control for the extracts throughout the period. Germination was completed
on day 18 at 92% for the control, day 20 at 95% for cineole and day 19 at 89% for
camphor.

For A. littoralis (Appendix 1, Figure A1.20) the germination occurred simultaneously


for all treatments on day 5 at 23% for the control, 28% for cineole and 29% for
camphor. Similar germination occurred throughout the period for all treatments with
only slight deviations above and below the control for the extracts applied.
Completion of germination occurred on day 26 at 46% for the control, day 28 at
44% for cineole and day 26 at 45% for camphor. Stepped germination also
occurred in 3 episodes which were displayed in all 3 treatments.

In the case of *C. coluteoides (Appendix 1, Figure A1.21) germination started at day
4 for cineole at 1% and day 6 for both camphor and the control at 11% and 15%
respectively. Overall germination in the treatments was similar to the control
throughout the trial. However, a slight delay occurred for cineole until day 10 when
it returned to the control level, and for camphor a slight delay with a fluctuation
slightly above the control at day 7 to 10, then returned to near control levels.
Germination was completed on day 10 for the control at 36%, cineole on day 13 at
38% and day 10 at 38% for camphor.

For the species A. cunninghamii (Appendix 1, Figure A1.22) germination commenced


on day 6 for all treatments, with 4% for the control, 8% for cineole and 4% for
camphor. Overall germination of all treatments was very similar throughout the
germination period with the 3 stepped germination sequences being similar in time
between each of the treatments. Germination was completed in the control on day 20
at 32%, day 18 at 33% for cineole and day 18 at 32% for camphor.

Germination of T. laurina (Appendix 1, Figure A1.23) commenced on day 8 for the


control at 2% and day 11 for both the cineole and camphor treatments both at
9%. Throughout the period, germination levels were similar in all treatments with
84

both the extract treatments being only slightly lower than the control. However,
germination continued until day 46 at 20% for the control while both the extracts
had completed germination at day 17 with cineole at 17% and camphor at 14%. A
stepped sequence of 4 episodes in the germination also occurred, but was most
pronounced in the control.

With F. coronata (Appendix 1, Figure A1.24) germination started in the control on


day 11 at 1% while for the extract treatments it commenced on day 15 for cineole at
6% and day 21 for camphor at 3%. Overall germination rates were lower than the
control initially for both the extracts, with camphor having the most pronounced
delay. Both the extracts then exceeded the germination levels of the control at day
19 for cineole and day 32 for camphor for the remainder of the germination.
Germination was completed for the control on day 40 at 69%, cineole on day 36 at
98% and camphor on day 54 at 87%. Stepped germination was also displayed for
this species with 4 episodes in all treatments which were in most cases simultaneous.

Germination of C. bartramia (Appendix 1, Figure A1.25) commenced on day 8 for


the control at 1%, day 16 for cineole at 1% and day 18 for camphor at 1%. Both
extracts displayed a pronounced delay in germination rate with the camphor being
substantially lower throughout the germination period, while cineole exceeded the
control levels at day 43 and slowly increased above the control from that point.
Germination was completed earlier for the control on day 46 at 72%, day 64 at 83%
for cineole and day 59 at 40% germination for camphor. Stepped germination was
also displayed for this species with 6 episodes identified in the control and a less
pronounced stepping in the extract treatments.

A. melanoxylon (Appendix 1, Figure A1.26) started germination on day 6 at 1% for


the control, day 10 for cineole at 5% and day 7 for camphor at 1%. Overall
germination levels were initially below the control until day 14 for camphor, and
day 27 for cineole, when they exceeded the control for the remainder of the species
trial. The stimulation above the control was also substantially greater for the
camphor than the cineole. Completion of germination occurred on day 30 at 50%
for the control, day 33 at 56% for cineole and day 29 at 72% for camphor.

85

The species E. microcorys (Appendix 1, Figure A1.27) commenced germination on


day 7 for the control at 2%, day 10 for cineole at 1% and day 8 for camphor at
2%. The germination rate remained lower than the control for both extracts initially,
with cineole being slightly less than the camphor. At day 21 the cineole then
exceeded the control and increased slightly for the remainder of the germination
period while the camphor approached a similar level to the control at day 16,
remaining slightly below that level. Germination was completed on day 28 at 29% for
the control, day 30 at 34% for cineole and day 16 at 26% for camphor.

For *L. camara (Appendix 1, Figure A1.28) a most pronounced delay in the start of
germination was exhibited in all treatments with the control and camphor
commencing on day 108 at 1% while for cineole no germination occurred during the
145 days of the species trial. For the stimulation shown in the camphor treatment,
germination steadily increased above the control at day 120. Germination was not
fully completed for this species at the termination of the trial due to the lengthy
dormancy period of the seed. However, at day 124, germination was 2% for the
control, and at day 145 it was 0% for the cineole and 6% for camphor.

In C. rubra (Appendix 1, Figure A1.29) very low germination occurred being only
1% in the control starting and completing on day 18, while in the treatments it was
0%. Seed viability may have been an issue with this species.

The germination of A. excelsa (Appendix 1, Figure A1.30) was also low being
recorded at 0% for all treatments during the trial of that species due to dormancy
which had not been released through hot water scarification treatment.

With D. pruriens (Appendix 1, Figure A1.31) germination started on day 13 at 5% for


the control, day 14 at 2% for cineole and day 14 at 10% for camphor. However,
rat predation occurred in the cineole resulting in the total loss of replication 1 and
2 i.e. half the cineole treatment. Overall, germination following this predation
resulted in similar germination rates for both the control and camphor, with the
camphor exceeding the control on day 19, gradually increasing beyond that point.
Germination was completed on day 22 at 94% for the control and day 27 at 103%
for the camphor, having poly-embryonic germination. However, in the undamaged
86

replication 3 and 4 of the cineole treatment, germination was completed on day 27


at 50% which is similar to the camphor.

In the species M. azedarach (Appendix 1, Figure A1.32) germination commenced on


day 14 for cineole at 1%, day 17 at 2% for the control and day 18 at 4% for
camphor. In the control, excess of TN and TP was detected at day 33 when
germination was at 22% with data recording ceasing at that time as a result of error.
However, in the extract treatments cineole reached higher levels of germination than
the camphor for most of the trial. At day 70 the camphor slightly exceeded the
cineole with both treatments still actively germinating. The germination was
completed on day 81 for both extract treatments with camphor at 95% and cineole
at 93%.

The germination of *C. camphora (Appendix 1, Figure A1.33) from fresh depulped
seed resulted in an error in the control when desiccation of seed was noted. This
prevented the control from attaining higher germination than 3% and indicated the
sensitivity of the seed to desiccation events. Other species germinating in the
propagation units of the control were unaffected by moisture stress. This occurred
over a 2 day period during high temperatures at day 28 of that species trial. However,
both the extract treatments remained unaffected, with camphor at first slightly
exceeding the cineole until day 80 when the cineole increased above that level for
the remainder of the germination period. In the camphor, germination was
completed on day 172 at 86% while in the cineole it occurred on day 168 at 89%. A
dormancy period of 27 days was exhibited for this species in the extracts with a
further slow germination period of 34 days from day 27 to day 61 when germination
only increased to 2% in the camphor. Following this delay, germination increased
rapidly and asynchronously for a lengthy period of 115 days.

F. xanthoxyla (Appendix 1, Figure A1.34) germination also showed an error in the


control with TN and TP levels in excess. The germination occurred simultaneously
in all treatments on day 14 at 5% for the control and cineole, and 1% for
camphor. Recording ceased for the control on day 21 when germination was at
23%. For the extract treatments which were unaffected by the error, cineole initially
germinated at a slightly higher rate than camphor which then exceeded it at day 18
87

until day 34 when it fell below the cineole level slightly. Germination was
completed on day 49 at 90% for the cineole and day 34 at 84% for the camphor.

For C. australe (Appendix 1, Figure A1.35) excess of TN and TP was also discovered
in the control. However, examination of the graph identified that germination may
not have been grossly affected due to the seeds very large cotyledon reserves. The
control and the camphor started germination on day 14 both at 1%, while the
cineole commenced on day 15 also at 1%. At first the germination of both the
extracts was slightly less than the control until day 21 when the cineole exceeded
the other treatment slightly for the remainder of the trial. The cineole exceeded the
control on day 23 and remained at a similar level until recording ceased in the
control on day 40. The germination was completed for the camphor on day 61 at
74% and for the cineole on day 67 at 86%.

In J. pseudorhus (Appendix 1, Figure A1.36) germination was noted to be in error for


the control due to further TN and TP excess. Germination started in the cineole
and camphor on day 16 at 2% and 3% respectively while the control commenced
on day 17 at 1%. All treatments remained at similar levels until day 22 when the
camphor exceeded the other treatments for the remainder of the trial. The control
however, remained between the 2 extract treatments until day 40 at 35% when
recording was terminated due to the error. The cineole completed germination on
day 55 at 37% and the camphor on day 53 at 49%.

D. australis (Appendix 1, Figure A1.37) germination also displayed an error in the


control where very low germination of 2% occurred resulting from water-logging
when punnets were positioned on sand which quickly became saturated once detritus
and algae impeded drainage. However, the error did not occur in the extract
treatments which both had similar germination until day 16 when the cineole
become slightly higher than the control. The germination was completed on day 40
at 29% for cineole, day 47 at 26% for camphor and day 37 at 2% for the control.

In the vines and climbers, *C. grandiflorum (Appendix 2, Figure A2.1) germination
started on day 6 for all treatments with the control and cineole at 1% and
camphor at 2%. Overall germination levels of the extracts treatments steadily
88

exceeded the control at a similar level until day 13 when the camphor exceeded the
level of the cineole for the remainder of the germination. The control however
gradually increased until it was at an equivalent level to the cineole on day 18.
Germination was completed for all treatment on day 22 at 61% germination for the
control, 59% for cineole and 76% for the camphor.

With the strike of *A. cordifolia tubers, (Appendix 2, Figure A2.2) the control and
cineole started shoot growth at day 15 at 6% and 5% respectively, while the
camphor started at day 12 at 2%. The overall germination rate of both the extract
treatments remained below the control from day 16 onwards, with each treatment at
a similar level although some fluctuation occurred. The strike was completed for all
treatments on day 94 at 75% for the control, 64% for cineole and 69% for
camphor. A stepped strike also occurred in this species with 8 episodes in the
control, 4 episodes in the cineole and 8 episodes in the camphor.

C. hypoglauca (Appendix 2, Figure A2.3) displayed a pronounced dormancy of 42


days before germination commenced in the control at 1%. In the cineole treatment,
germination started on day 50 at 1% and in the camphor at day 48 at 1%. The
germination rate of both extracts was lower than the control initially, with the
cineole remaining lower throughout the germination, while the camphor exceeded
the control at day 98 and steadily increased. Completion of germination occurred
simultaneously for all treatments at day 122 when the control was at 73%, cineole
at 65% and the camphor at 83% germination. Stepped germination also occurred for
this species with as many 6 episodes in the control, 8 episodes in the cineole and 9
episodes in the camphor.

The germination of G. cymosum (Appendix 2, Figure A2.4) showed a pronounced


dormancy of 60 days before germination commenced in the control at 1%. However,
this delay was longer for the cineole being 102 days at 1%, and the camphor with
no germination during the period of the species trial. However, the cineole increased
above the level of the control at day 126 when germination reached the highest level
of 7%. With the control, germination reached maximum for that treatment at day 91
being only 2%. The trial was terminated at day 141 for this species, and it is presumed

89

that germination rates would have increased over a further period of time. Seed
viability may have also influenced the germination rate of this species.

For *P. suberosa (Appendix 2, Figure A2.5) germination was low with 2%
germination for camphor at day 20 and 1% for cineole also at day 20. The
control remained at 0% germination for the entire period of the trial. This suggested
that seed viability may have been an issue.

In the herbs, O. aemulus (Appendix 3, Figure A3.1) started germination on day 10 in


the control at 1% and day 13 for both cineole and camphor at 3% and 12%
respectively. The germination rate increased above the control for both extract
treatments throughout the germination period, with the camphor maintaining the
highest germination level. Germination was completed on day 60 at 80% for the
control, day 57 at 90% for cineole and day 60 at 97% germination for camphor.
A stepped germination was also noted with 4 sequences for the control and
cineole, while the camphor had 5 sequences.

For C. enervi, (Appendix 3, Figure A3.2) germination commenced on day 18 at 1%


for the control, day 21 at 2% for cineole and day 26 at 1% for camphor. Both
extracts were slightly stimulated initially, with the cineole then returning to similar
levels to the control, while the camphor gradually increased above the control for
the duration of germination. The germination was completed in the control on day
86 at 13%, day 124 at 14% for cineole and day 133 at 30% for camphor. A
sequence of germination stepping was also shown in this species, with 4 episodes in
the control, 8 episodes in the cineole and 12 episodes in the camphor.

*S. sphacelata var. narok (Appendix 3, Figure A3.3) germination started on day 7 for
all treatments with the control at 4%, cineole at 8% and camphor at 5%. The
overall germination rates were similar for the control and camphor throughout the
germination, while cineole was stimulated above the control. Germination was
completed for the control on day 15 at 12%, for cineole on day 22 at 22%, and day
20 at 14% for camphor.

90

In the species O. gracillima (Appendix 3, Figure A3.4) germination commenced on


day 8 at 4% for the control, day 7 at 1% for cineole and day 10 at 3% for
camphor. Initially, germination in the camphor treatment was slightly below the
control but at day 17 this became slightly higher until day 55 when the level of the
control matched that of the camphor. In the cineole however, germination
exceeded the control slightly from day 10 to day 33 when the control approached a
similar level. Completion of germination occurred on day 55 at 16% for the control,
day 53 at 17% for cineole and day 27 at 16% for camphor.

The germination of A. caerulea (Appendix 3, Figure A3.5) started on day 31 at 1% for


the control, day 47 at 3% for the cineole and day 41 at 3% for camphor,
indicating a long dormancy period. The germination was similar for the control and
camphor until day 73 when the camphor fluctuated slightly above the control
then fell below it on day 99, where it remained. The cineole however, remained
below both the germination levels of the other treatments, germinating slowly until
day 139 when it remained at 49% germination. For the control, germination was
completed on day 127 at 52% and for the camphor on day 135 at 12%.

*C. gayana (Appendix 3, Figure A3.6) commenced germination on day 3 for all
treatments, at a rate of 12% for the control, 10% for cineole and 3% for camphor.
Although initial germination of the cineole was slightly below the control this
treatment become similar to the control on day 5 and remained just below the
control for the remainder of the germination. The camphor germination however,
remained below the other treatments for the entire period of the trial. Germination was
completed on day 9 at 42% for the control, day 16 at 41% for cineole and day 25 at
39% for camphor.

For V. hederacea (Appendix 3, Figure A3.7) cutting strike started on day 4 at 1% for
the control, and day 7 for both cineole and camphor at 7% and 12% respectively.
Strike rates remained below the level of the control for both these treatments with
cineole being slightly lower than the camphor throughout the strike period.
Treatments completed the strike on day 25 at 98% for the control, and day 19 for
both cineole and camphor at 72 and 77% respectively.

91

In the species *P. clandestinum (Appendix 3, Figure A3.8) germination developed


rapidly, starting in the control on day 4 at 1%, while on day 5 it increased markedly
to 77%. For the cineole and camphor germination started on day 5 at 5% and 1%
respectively, and substantial increases to 36% in cineole and 43% in camphor
occurred on day 6. The overall germination rates for cineole and camphor were
maintained below the level of the control for the entire germination period, with the
cineole being slightly below that of the camphor. Completion of germination
occurred on day 12 at 96% for the control, day 18 at 84% for cineole and day 21 at
87% for camphor.

The cutting strike of P. purpurascens (Appendix 3, Figure A3.9) started on day 5 for
all treatments with the control at 9% and cineole and camphor both at 17%. Both
treatments remained above the level of the control until day 24 when the cineole
became similar to the control, and day 15 for camphor, when these levels dropped
below the control for the remainder of the strike period. Strike of this species was
complete for both the control and cineole on day 28 at 91% and 92% respectively,
while for the camphor it as completed on day 33 at 78%.

For *P. dilatatum (Appendix 3, Figure A3.10) germination commenced on day 8 at


1% for the control, day 10 at 1% for cineole and day 21 at 1% for camphor. The
overall germination rate for both the extract treatments remained below the control
level for the entire germination, with camphor being substantially lower than the
cineole. For the control, germination was completed on day 48 at 56%, while for
the cineole and camphor it was day 55 at 54% and 9% respectively. A stepped
germination was also demonstrated in this species, with 3 short episodes in the
control, 4 episodes in the cineole and 2 episodes in the camphor.

H. bracteatum (Appendix 3, Figure A3.11) germination started on day 6 for the


control and cineole at 7% and 1% respectively, while the camphor was 1% at day
9. The level of germination in the cineole remained slightly below the control for
the entire germination period. The camphor at first was slightly lower than the
cineole, then at day 5 it exceeded the camphor and slowly increased until it
became equivalent to the control at day 57. Completion of germination occurred on

92

day 41 at 15% for the control, day 47 at 10% for cineole and day 57 at 15% for
camphor.

With L. longifolia (Appendix 3, Figure A3.12) germination commenced on day 28 at


1% for the control, and day 34 at 1% for both cineole and camphor. Germination
levels were fairly similar initially for the control and cineole, with a minor delay in
the cineole until day 43 when levels became almost equivalent to slightly below the
control. However, on day 73 the cineole exceeded the control and increased
substantially for the remainder of the germination. In camphor, germination levels
remained below both the other treatments for a lengthy period until day 88 when it
exceeded the control, and further increased until it become equivalent to the
germination levels of the cineole on day 104. Germination was completed for this
species on day 101 at 62% for the control, day 101 at 81% for cineole and day 108
at 83% for camphor. A stepped germination was also apparent with the control,
displaying 5 episodes, the cineole with 6 episodes and the camphor 5 episodes.

A comparison of the number and percentage of species which display variability in


the commencement and completion time of germination compared with the control
is summarized in Figure 9a and b respectively. The graphs identify that both camphor
laurel chemo-types produce relatively similar responses to the extracts in both
germination commencement and completion. A substantial delay for many species in
both commencement and completion time was identified while a smaller number
exhibited early germination commencement and completion. Overall 68% or 30 of the
44 vascular species displayed a similar response to the extract treatments for
germination commencement as well as the completion. However, only 16% or 7 of
the 44 species had the same response between germination commencement and
completion.

The species germination responses to the individual extracts when compared with
control fall into 7 groups, with species most often responding differentially to the
cineole and camphor in their rate of germination. This comparison included 44
species excluding errors and germinations below 5% in 10 species. The analysis
includes both slight to substantial responses and are pre-statistical. The effect of the
individual extracts in these response groups is provided in Table 9. The table also
93

Table 9. The distinct germination response groups, species (a) and seed
characteristics (b) influenced by the cineole and camphor treatments.
(a). Germination Response Group and Species

Response Group

Response 1:
(stimulation followed
by a return to control)

Response 2:
(stimulation followed
by a delay below the
control)

Response 3:
(full stimulation above
the control)

Response 4:
(delay followed by a
return to the control)

Response 5:
(delay followed by
stimulation above the
control)

Response 6:
(full delay below the
control)

Species for
Cineole

Species for
Camphor

S. paniculatum
C. enervis
O. gracillima
P. purpurascens
*C. grandiflorum

M. tanarius
M. philippensis
A. caerulea

Myrtaceae (1)
Poaceae (1)
Lobeliaceae (1)
Sapindaceae (1)
Euphorbiaceae (2)
Zingiberaceae (1)
Cyperaceae (1)

Trees (3)
Herbs (4)
Vines (1)

T. australis
L. confertus
L. longifolia

T. australis
S. paniculatum
L. longifolia
P. purpurascens

Meliaceae (1)
Myrtaceae (2)
Xanthorrhoeaceae (1)
Lobeliaceae (1)

Trees (3)
Herbs (2)
Vines (0)

M. philippensis
F. macrophylla
O. aemulus
*S. sphacelata var. narok

E. intermedia
*L. camara
O. aemulus
C. enervis
*C. grandiflorum
G. semiglauca

Euphorbiaceae (1)
Moraceae (1)
Poaceae (2)
Myrtaceae (1)
Verbenaceae (1)
Sapindaceae (2)
Cyperaceae (1)

Trees/
Shrubs (5)
Herbs (3)
Vines (1)

E. intermedia
G. semiglauca
S. leuhmannii
O. populifolius
*C. gayana
*P. dilatatum

H. bracteatum
*L. lucidum
S. luehmannii
O. populifolius
*C. gayana
H. flavum
E. microcorys
*O. gracillima

Myrtaceae (3)
Sapindaceae (1)
Euphorbiaceae (1)
Poaceae (3)
Asteraceae (1)
Oleaceae (1)
Pittosporaceae (1)

Trees (7)
Herbs (4)
Vines (0)

F. coronata
C. bartramia
A. melanoxylon
E. microcorys
*P. clandestinum
Cissus hypoglauca

F. coronata
L. pulchella
A. melanoxylon
E. saligna
*P. clandestinum
G. cymosum

Moraceae (1)
Sterculiacea (1)
Mimosaceae (1)
Myrtaceae (2)
Poaceae (1)
Vitaceae (1)
Sapindaceae (1)
Philesiaceae (1)

Trees (6)
Herbs (1)
Vines (2)

E. saligna
*A. cordifolia
M. tanarius
V. hederacea
*L. lucidum
*S. capsicoides
*C. camphora (from soil)
A. torulosa
*L. camara
A. caerulea
H. bracteatum
G. cymosum

L. confertus
*A. cordifolia
C. bartramia
V. hederacea
*P. dilatatum
*S. capsicoides
*C. camphora (from soil)
A. torulosa
C. hypoglauca

Sapindaceae (1)
Myrtaceae (3)
Euphorbiaceae (1)
Oleaceae (1)
Solanaceae (1)
Pittosporaceae (1)
Lauraceae (1)
Casuarinaceae (1)
Verbenaceae (1)
Sterculiaceae (1)
Basellaceae (1)
Philesiaceae (1)

Trees/
Shrubs (12)
Herbs (4)
Vines (3)

94

Families

Lifeforms

No. of
Species
Influenced
by Both
Treatments

Vitaceae (1)
Zingiberaceae (1)
Violaceae (1)
Asteraceae (1)
Poaceae (1)

H. flavum
C. viminalis
L. pulchella

Response 7:
(response similar to
the control)

C. parvifolia
A. littoralis
*C. coluteoides
A. cunninghamii
T. laurina

C. parvifolia
A. littoralis
*C. coluteoides
A. cunninghamii
T. laurina
*S. sphacelata var. narok
F. macrophylla
C. viminalis

Sapindaceae (1)
Casuarinaceae (1)
Caesalpinaceae (1)
Araucariaceae (1)
Myrtaceae (2)
Moraceae (1)
Poaceae (1)

Trees/
Shrubs(7)
Herbs (1)
Vines (0)

(b). Germination Response Group, Seed Characteristics and Dormancy

Response Group

Seed Weight
Range of 100 seeds
(grams)

Ratio
Dry to
Moist
Seed

No. of
Introduced
Species

Ratio Dormancy
to No Dormancy

Response 1:

0.01 12.08

4:5

1:7

0.02 - 11

3:2

0:5

0.01 12.08

6:3

2:7

0.013 -12.77

7:4

0:11

0.02 -7.79

6:3

5:4

0.008 - 52.52

10:9

9:10

0.008 30.33

6:2

0:7

(stimulation followed
by a return to control)

Response 2:
(stimulation followed
by a delay below the control)

Response 3:
(full stimulation above the control)

Response 4:
(delay followed by a return to the
control)

Response 5:
(delay followed by stimulation above
the control)

Response 6:
(full delay below the control)

Response 7:
(response similar to the control)

95

identifies the species reacting in each response group to the cineole or camphor
treatment, the number of species represented in the families, the number of life-form
groups of the species, the seed weight range for all species, the ratio of dry to moist
seeds, the number of introduced species, the ratio of dormancy to no dormancy of the
seed and the number of species influenced by both treatments.

No. of species

(a).

45
40
35
30
25
20
15
10
5
0

n = 44

early
same
delay

45%
48%
36%

39%

%
16%

16%

Cineole

Camphor

Treatment and No. of species exhibiting similar responses to both leachates.

(b).

Figure 9. Number and percentage of variability in: (a) commencement


of germination and (b) completion of germination compared to the
control.

Figure 10 provides the number and percentage of species in the sample which respond
to the 7 groups through a comparison of the individual extract treatments against the
control. Sample n in the life-form groupings is higher than species n due to the
96

Germination response groups:


34%

16

cineole

No.of species

14
12

camphor
n =44

10

20%

18%

14%

6 11%
7%

14%

18%

14% 14%
11%

9%

9%

7%

2
0
Group 1:
Group 2:
Stim/Return Stim/Delay

Group 3:
Group 4:
Group 5:
Full Stim Delay/Return Delay/Stim

Group 6:
Full Delay

Group 7:
Similar to
Control

Response group

Stim/Return
Stimulation above control then return
to control.
Stim/Delay
Stimulation above control then delay
below control.
Full Stim
Maintained stimulation above control.
Delay/Return
Initial germination below control then
a return to control.
Delay/Stim
Initial germination below control then
stimulation above control.
Full Delay
Maintained delay in germination below
control.
Similar to Control
Germination at similar level to
control.

Figure X. Number and percentage of species in various response


Figure
10. Number and percentage of species in various response
groups compared against the 'control' for the 'cineole' and
groups compared
against the control for the cineole and camphor
'camphor' treatments.

treatments.

differential effect of the extracts, which means that a species can occur in two
response groups. This also applies to other graphs in this summary.

The number of vascular plant families in each response group is compared in Figure
11. This graph also identifies the distribution of the families in each response group
which provides a further comparison of any family relationships and number of

No. of families

species trialed in each family.

18
16
14
12
10
8
6
4
2
0

17

n of species = 44
n of families = 26

Group 1:
Group 2:
Stim/Return Stim/Delay

Group 3: Group 4:
Group 5:
Full Stim Delay/Return Delay/Stim

Group 6:
Full Delay

Response group

Group 7:
Similar to
Control

Family
Group Location
Myrtaceae
all
Poaceae
1,3,4,5,6,7
Cyperaceae
1
Lobeliaceae
1,2
Sapindaceae
1,3,4,5,6,7
Euphorbiaceae
1,3,4,5,6,7
Zingiberaceae
1,6
Meliaceae
2
Xanthorrhoeaceae 2
Moraceae
3,5,7
Verbenaceae
3,6
Cyperaceae
3
Asteraceae
4,6
Oleaeceae
4,6
Pittosporaceae
4,6
Sterculiaceae
5,6
Mimosaceae
5
Vitaceae
5,6
Philesiaceae
5,6
Solanaceae
6
Lauraceae
6
Casuarinaceae
6,7
Basellaceae
6
Violaceae
6
Caesalpinaceae
7
Araucariaceae
7

Note: For germination


response groups refer
to Figure 10.

Number
of plant
families
various response groups
Figure
Number
of plant families
in each
responsein
group.
Figure X.11.
compared against the control for the cineole and camphor
treatments.
97

A comparison of the life-form groupings which include vines and twiners, herbs and
trees and shrubs, is provided in Appendix 4, Figure A4.1.

Relationships involving the weight of seeds, cuttings, tubers or stolons within and
between each of the response groups are compared in Appendix 4, Figure A4.2.

A summary of the number of dry and moist propagules in the response groups is
shown in Appendix 4, Figure A4.3.

The proportion of native plants to introduced species in each of the response groups is
compared in Appendix 4, Figure A4.4.

Species with similar and distinct responses to the two extracts in the response groups
is shown in Appendix 4, Figure A4.5, and dormancy is compared to rapidly
germinating species in Appendix 4, Figure A4.6.

2.4.5 Soil Algae Trialed and Pre-Statistical Results of Growth


The soil algae identified from the cryptogamic mat of the first algal trial are shown in
Table 10. This included 27 species, 20 in the Division Cyanobacteria, 5 in
Chlorophyta and 2 in Bacillariophyta. The filamentous growth form included 14
species, while uni-cellular algae numbered 12 species, and multi-cellular algae
without filamentous habit, 1 species.

Results from the collective growth of soil algae obtained from three locations are
shown in Figure 12 a, b and c.

In sample 1, Figure 12a, the control commenced visual growth on day 4 with full
coverage of the punnets in 20 days. For cineole, observed growth commenced on
day 34 covering the punnets on day 52, while for camphor growth begun on day 57
with coverage on day 100.

For sample 2, Figure 12b, algae in the control commenced visual growth at day 9
and covered the media surface by day 16. The treatments also showed slower

98

Table 10. Soil algae identified in the first algal trial through sampling of the media and
microscopic examination.
Cyanobacteria
Anabaena
Calothrix
Closteridium
Cylindrospermum
Entophysalis sp.1
(Hydrococcus)

Entophysalis sp.2
(Hydrococcus)

Gloechaete?
Haematococcus
Microchaete
Nodularia
Nostoc
Phormidium
Pseudoanabaena
Pseudotetraspora
Spirulina
Stigonema
Unknown sp. No. 17
Unknown sp. No. 20
Unknown sp. No. 21
Unknown sp. No. 5

Cell
Organization

Chlorophyta

Cell
Organization

filamentous
uni-cellular
uni-cellular
filamentous
shortfilaments
shortfilaments
uni-cellular
uni-cellular
filamentous
filamentous
filamentous
filamentous
filamentous
uni-cellular
uni-cellular
filamentous
uni-cellular
uni-cellular
uni-cellular
uni-cellular

Cylindrocapsa
Oedocladium sp.1
Oedocladium sp.2
Pamellopsis
Ulothrix

filamentous
filamentous
filamentous
multi-cellular
filamentous

Bacillariophyta

Cell
Organization

Amphora
Peronia

uni-cellular
uni-cellular

responses with cineole starting growth at 21 days and completing at 42 days , and
camphor starting at 40 days, covering the punnet at 97 days.

Growth of algae for sample 3, Figure 12c, also displayed similar response to the first
two samples. In the control, algae started growing at day 8 and completed coverage
by day 17. For cineole, growth began on day 25 and covered the punnet by day 66.
Similarly, camphor which also showed the slowest response, started at day 55 and
completed coverage of the media at day 100.
For sample 1 only, abundance and diversity of all species in each treatment is
displayed in Figure 13. There is a reduction in species diversity from 23 species in the
control to 9 species in the camphor and 12 species in the cineole treatments i.e. a
reduction in species diversity of 60% and 38% respectively. Mean abundance of all
algae however, reduced slightly with only small reductions in the treatments
compared to the control i.e. reducing from an abundance score of 1.34 in the
control to 1.16 for camphor and 1.27 for cineole, providing a 13% and 5%
reduction in mean abundance respectively.

99

(a).
100
90
80
70

% Cover

60
50
40
30

C o n t ro l
C in e o le
C am phor

20
10
0
1

7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 76 79 82 85 88 91 94 97

D a yDay
1.
F ig u re X a . G ro w th o f a lg a e fro m s a m ple

(b).
10 0
90
80

% Cover

70
60
50
40
30

C on tro l
C in eo le
C am p h or

20
10
0
1

7 10 1 3 16 19 2 2 25 2 8 3 1 34 3 7 40 43 4 6 49 5 2 5 5 58 6 1 64 67 7 0 73 7 6 7 9 82 8 5 8 8 9 1 9 4 97

D ay

2.
F ig u re X b . G ro w th o f a lga e fro m s a mpleDay

(c).
100
90
80

% Cover

70
60
50
40
Control

30
20

Cineole

10

Camphor

0
1

7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 76 79 82 85 88 91 94 97

Day
Day

Figure 12. Growth of algae over the media surface expressed as


percentage cover over time for: (a) sample 1; (b) sample 2; (c) sample 3.

100

25

23

Diversity
Mean Abundance

20
Species Diversity
& Abundance
score

15

12

Treatment:

10
1.16

1.34

1.27

Co - control
Ca - camphor
Ci - cineole

0
Co

Ca

Ci

Treatment

Diversity and
and mean abundance
Figure.
abundanceofofalgae
algae
Figure
13. XDiversity
the treatments
algae sample
1.
across theacross
treatments
in algaeinsample
1.

In sample 1, the 27 species recorded are represented in 5 distinct response groups to


the extracts:

Group 1: a high abundance in the control and decreased abundance in the


treatments which included Cyanobacteria sp.5, Pseudanabaena, Nodularia,
Pamellopsis, Pseudotetraspora and Cylindrocapsa, Figure 14;
Group 2: high abundance in the control and differential abundance in the
treatments with only Nostoc represented, Figure 15;
Group 3: low abundance in the control and absent in the treatments for the
species Ulothrix, Oedocladium sp.1, Oedocladium sp.2, Haematococcus,
Entophysalis sp.1, Gloechaete?, Cyanobacteria sp. 21, Microchaete,
Cylindrospermum and Closteridium, Figure 16;
Group 4: absent or low abundance in the control and increased abundance in
the treatments with Calothrix, Phormidium, Amphora and Anabaena, Figure
17; and
Group 5: low abundance in the control and differential abundance in the
treatments which included Stigonema, Peronia and Cyanobacteria sp.17,
Figure 18.

101

co mmo n

3
2.8
C o - C o ntro l
C a - C am pho r
C i - C ineo le

2.6
2.4
2.2
A bunda nc e
s c o re

2
1.8
1.6
s p5. C yano bac t eria
s p7. P s eudanabaena
s p2. N o dularia
S pe c ie s
s p1. P am ello ps is
s p22. P s eudo tet ras po ra
C lo s teridium
s p6. C ylindro c aps a

1.4
1.2
ab sent

Co

Ca

Ci

T re a t m e n t

Figur e . Xa Sample1 algae of 'high relative ab undanc e' in the

Sample
1reas
algae
of high
relative
abundance
Figure 14.
ed relative
ab undanc
e'' in the
control
and 'dec
treatments . in
the control and decreased relative abundance in the
camphor and cineole treatments.
co mmo n

3
2 .8

C o - C o n t ro l
C a - C am pho r
C i - C in e o le

2 .6
2 .4
A bundance
s c o re

2 .2
2
1.8
1.6
1.4

ab s ent

1.2
1

Co

s p 10 . N o s t o c

Ca

Ci

T re a tm e nt

S p e c ie s

g u r e . X b S a m p le 1 a lg a e o f 'h ig h re la t iv e a b u n d a n c e ' in
Sample 1 algae of high relative abundance in the
FigureFt hi15.
e c o n t ro l a n d 'd if f e re n t ia l a b u n d a n c e ' in t h e t re a t m e n t s .
control and differential abundance in the camphor and
cineole treatments (Nostoc only).

102

co mmo n

C o - C o ntro l
C a - C am pho r
C i - C ineo le

3
2 .8
2 .6
2 .4

A bunda nc e
s c o re

ab sent

2 .2
2

sp3. Cy lindr osper m um


sp.24 M ic r oc haet e
sp.21 Cy anobac t r eia
S p e c ie s
sp.15 Gloec haet e ?
sp.14 Ent ophy salis sp.1
sp. 13 Haemat oc oc c us
sp.9 Oedoc ladium sp.2
sp.8 Oedoc ladium sp.1
sp.4 Ulot hr ix

1.8
1.6
1.4
1.2
1

Co

Ca

Ci

T re a t m e nt

1 algae of low relative abundance in


Figure 16. Sample
Figur e . Xc Sample 1 algae of 'low relative ab undanc e' in the
the control and absentcontrol
in theand
camphor
cineole
treatments.
'ab s ent'and
in the
treatments.

C o - Co nt ro l
C a - C amp ho r
C i - Cineo le

co mmo n

2.8
2.6
2.4
2.2

A bunda nc e
s c o re

2
1. 8

sp 2 5 A nab aena

1.6

sp .2 6 A mp ho ra

1.4

ab sent

sp .12 Pho rmid ium

1. 2

S p e c ie s

sp .18 Sp irulina

Co
T re a t m e nt

sp .2 7 Calo t hrix

Ca

Ci

Figure Xd. Sam ple 1 algae either 'ab sent' or with 'low
Figure
17. Sample 1 algae either absent or with low relative
relative ab undance' in the control and 'increased relative
abundance in the
control and increased relative abundance in the
ab undance' in the treatm ents .
camphor and cineole treatments.

co mmo n

3
2.8
2.6

C o - C o ntro l
C a - C am pho r
C i - C ineo le

2.4

A bunda nc e
s c o re

2.2
2
1.8
1.6
1.4

s p.17 C yano bac teria


s p. 20 C yano bac teria
s p.19 P ero nia
s p.23 Ento phys alis sp.2
S p e c ie s
s p.11 Stigo nem a

1.2
ab sent

Co
T re a t m e n t

Ca

Ci

1 algae
of 'low
relativeabundance
ab undance'
Figure.
Xe Sam ple
1 algae
of low
relative
in in
Figure
18. Sample
the control and 'differential' effect in the treatm ents
the control and differential abundance in the camphor and
cineole treatments.

103

2.4.6 Seed Germination and Algal Growth Significant Effects of the


Treatments
Non-linear regressions performed on the germination profiles of each vascular species
and the algal growth profiles for the control, cineole and camphor treatments
using the Richards function appear in Appendix 5 for the trees and shrubs; Appendix
6 for the herbs, Appendix 7 for the vines and Appendix 13 for the algae. On average
the r2 of the Richards function exceeded 0.9 for most species identifying a close fit for
the modeled data to the actual growth curves. A summary of the modeled data from
the Richards function for the asymptote, mid-inflection point, days to midinflection, germination rate or growth coverage rate and the natural log of the
days to mid-inflection appear in Appendix 8.

From the deviance test, Appendix 9, the model estimated mean ( SE) for the
asymptote was (58.9 4.6), for the mid-inflection point (24.8 2.1), for the
germination or growth coverage rate (0.63 0.24), and for the natural log of the days
to mid-inflection (2.8 0.12).
2

The treatment effect was non-significant (p>0.05) for the asymptote (2 = 1.546, p =
2

0.462), mid-infection point (2 = 2.201, p = 0.333), and germination or growth


2

coverage rate parameters (2 = 2.525, p = 0.283). However, a highly significant


difference among treatments was identified for the natural log of the days to mid2

inflection point (2 = 12.505, p = 0.002).


The mean log of the days to mid-inflection at 95% CI was shortest in the control
treatment (2.44 - 2.69 - 2.94), longer in the cineole treatment (2.61 - 2.86 - 3.11),
and longest in the camphor treatment (2.68 - 2.93 - 3.18). The back-transformed
estimates for input to the multilevel analysis were for the control 14.79, the cineole
17.51 and the camphor 18.67. The multi-level analysis applied to these estimates at
95% CI were for the control (0.04 - 0.17 - 0.3), for cineole (0.1 - 0.23 - 0.36) and
for camphor (0.07 - 0.06 - 0.19). Planned comparisons between the pairs of
treatments found significant differences between both the cineole and control (p =
0.010) and the camphor and control (p = 0.0005), but the difference between the

104

camphor and cineole treatments was not significant (p = 0.335). This identified
that the cineole and the camphor treatments had greater effect than the control
while no significant difference existed between the effect of the cineole and
camphor treatments.

The residual plot of the asymptote, days to mid-inflection, mid-inflection point


and germination or growth coverage rate (Appendix 10) separates species of highest
treatment effect where the residuals occur above the mean (0) from the lowest effect
which occur below the mean (0). Confidence intervals for the data are also presented
on these residual plots providing a standard error for the germination data, not able to
be generated during the non-linear regressions using the Richards function in Excel
Solver.

For the residual plot of the asymptote for the vascular plants and algae, Appendix
10, Figure A10.1 the lowest effect ranged between -15.514 to -50.603 +/- 1.4 sd. For
vascular plants these were in descending order of effect: A. littoralis, *P. dilatatum,
*C. gayana, *C. colluteoides, A. caerulea, A. cunninghamii, L. confertus,

E.

microcorys, C. enervis, *C. camphora (from soil), T. laurina, O. gracillima, *S.


sphaecelata, H. bracteatum, A. torulosa, M. philippensis, F. macrophylla and *A.
cordifolia. Algae were not represented in this lower response group. The highest
effect on all the plants i.e. residuals occurring above the mean (0), ranged between
0.74894 to 45.711 +/- 1.4 sd. When arranged in ascending order of effect for the
vascular plants these included: H. flavum, A. melanoxylon, C. viminalis, *C.
grandiflorum, C. bartramia, M. tanarius, G. cymosum, S. capsicoides, C. hypoglauca,
E. saligna, G. semiglauca, L. longifolia, O. populifolius, V. hederacea, *P.
clandestinum, *L. lucidum, T. australis, F. coronata, S. paniculatum, P.
purpurascens, E. intermedia, S. luehmannii, L. pulchella, C. parvifolia, and O.
aemulus. In this effect group the asymptote of algae culture 3, algae culture 1 and
algae culture 2 respectively were found to be more highly affected than the seedling
asymptotes.

In the residuals for the natural log of the days to mid-inflection, Appendix 10, Figure
A10.2, the lowest treatment effect on the species with residuals below the mean (0)
varied between -0.01709 to -1.4537 +/- 1.4 sd.
105

Presented in decreasing order of

effect the vascular species included: F. macrophylla, O. populifolius, L. confertus, A.


melanoxylon, *C. grandiflorum, A. torulosa, S. paniculatum, O. gracillima, S.
luehmannii,

T. australis, E. microcorys, H. bracteatum, *P. clandestinum, V.

hederacea, C. parvifolia, T. laurina, L. pulchella, *S. sphaecelata, A. cunninghamii,


E. intermedia, P. purpurascens, G. semiglauca, E. saligna, *C. colluteoides, C.
viminalis, A. littoralis, and *C. gayana. Algae were not represented in this lower
response group. The highest influence of the treatments on species where the residuals
were above the mean (0) varied from 0.039869 to 1.6171 +/- 1.4 sd. Arranged in
order of ascending effect the vascular species included: M. tanarius, S. capsicoides, F.
coronata, C. bartramia, G. cymosum, O. aemulus, *C. camphora (from soil), *P.
dilatatum, C. enervis, *L. lucidum, A. caerulea, M. philippensis, L. longifolia, C.
hypoglauca, H. flavum and *A. cordifolia. The algal cultures were represented in this
higher effect group for the days to mid-inflection but did not show any dissimilarity
to how the seedlings responded as the three cultures were distributed through the midrange of the residuals.

For the mid-inflection point, Appendix 10, Figure A10.3, the lowest effect of the
treatments on species, where the residuals occurred below the mean (0), ranged
between -1.0197 to -20.569 +/- 1.4 sd. When arranged in descending order of effect
the vascular species represented were: A. melanoxylon, C. viminalis, *C. gayana, A.
littoralis, *C. colluteoides, A. caerulea, A. cunninghamii, L. confertus, E. microcorys,
*C. camphora (from soil), F. macrophylla, C. enervis, T. laurina, O. gracillima, *S.
sphaecelata, A. torulosa, H. bracteatum, M. philippensis and *Anredera cordifolia.
The algae were not present in this group. Species which occurred above the residual
mean (0) which demonstrated the highest effect of the treatments on species varied
between 0.57232 to 37.811 +/- 1.4 sd. Presented in ascending order of effect the
vascular species included: *P. dilatatum, M. tanarius, G. cymosum, C. hypoglauca,
G. semiglauca, E. saligna, H. flavum, C. bartramia, S. paniculatum, S. capsicoides, F.
coronata, P. purpurascens, L. pulchella, L. longifolia, S. luehmannii, C. parvifolia,
*C. grandiflorum, O. populifolius, E. intermedia, T. australis, *L. lucidum, V.
hederacea, *P. clandestinum, and O. aemulus. For this effect group the midinflection of algae culture 1, algae culture 2 and algae culture 3 respectively were
identified to be more highly affected than the seedlings except for O. aemulus which
fell midway between algae culture 1 and algae culture 2.
106

In the residual plot of germination or growth coverage rate (Appendix 10, Figure
A10.4) the lowest effect of the treatments on species identified below the residual
mean (0) was found to be -0.0040532 to -0.57289 +/- 1.4 sd. Presented in descending
order of effect these were: S. paniculatum, L. longifolia, A. melanoxylon, O.
gracillima, M. tanarius, L. confertus, H. bracteatum, G. cymosum, A. caerulea, C.
hypoglauca, M. philippensis, *L. lucidum, *P. dilatatum, C. enervis, H. flavum, *C.
gayana, F. macrophylla, O. aemulus, C. bartramia, *A. cordifolia and T. australis.
Algae culture 1, algae culture 3, algae culture 2 were found to be present in this
lowest effect range and due to their distribution throughout the seedling data
demonstrated similar response to the seedlings. The highest influence of the
treatments on species where the residuals were above the residual mean (0) varied
between 0.0003 to 1.1788 +/- 1.4 sd. Arranged in order of ascending effect the
vascular species included: V. hederacea, *P. clandestinum, F. coronata, *S.
sphaecelata, E. saligna, T. laurina, *C. camphora (from soil), Guioa semiglauca, S.
luehmannii, *C. grandiflorum, A. cunninghamii, L. pulchella, E. microcorys, C.
parvifolia, P. purpurascens, C. viminalis, S. capsicoides, A. torulosa, A. littoralis, E.
intermedia, *C. colluteoides and O. populifolius. Algae were not found to be present
in this group.

2.4.7 Vascular Plant Growth Significant Effects of the Treatments


The t-tests for the post-emergent measurements of shoot length, radicle length, leaf
number and leaf area are shown in Appendix 11. Summary effect tables generated
from the t-tests for each species appear in Appendix 12; Table A12.1 for the trees and
shrubs, Table A12.2 for the herbs and Table A12.3 for the vines.

When the sampled assemblage is considered as a whole i.e. trees, shrubs, vines and
herbs in combination, Figure 19a to d, the t-tests demonstrated that the significant
growth responses to both the applied extracts were 31 of the 36 species (86%) for
shoot length, 23 species (64%) for radicle length, 29 species (54%) for leaf number
and 25 species (71%) for leaf area.

The significant effect of the individual cineole and camphor treatments on shoot
growth of the sampled assemblage (Figure 19a) revealed that the cineole only effect
had 28% of species with significant reductions and 6% with significant increases in
107

growth while the camphor only effect had only 6% of species with significant
reductions and 11% with significant increases. A significant effect on shoot growth
from both extracts was also demonstrated with 25% of species having significant
reductions, 6% with significant increases and 6% with significant growth reductions
to cineole and a significant increase to camphor. No significant difference were
identified for 14% of species in the sampled assemblage.

For radicle length of the sampled assemblage (Figure 19b) the cineole only effect
contained 8% of species with significant growth reductions and 3% with a significant
increase while the camphor only had 17% significant reductions and 6% with
significant increases. The significant effect of both extracts on radicle length
accounted for 19% of species with significant reductions, 8% with significant
increases and 3% with a significant reduction to cineole and a significant increase to
camphor. The t-tests also showed that 36% of species had no significant growth
response to the extracts for radicle length.

In the significant effect of the treatments on leaf number of the sampled assemblage,
(Figure 19c), 11% of species were found to have significant growth reductions for
cineole only and 6% with significant increases while the camphor only had 6%
with significantly reduced growth. For the significant effect of both extracts 28% of
species had significant reductions in leaf number and 3% with a significant increase.
The treatments were also found to have no significant effect on the leaf number in
44% of species from the tested assemblage.

T-tests of leaf area of the sampled assemblage (Figure 19d) identified the significant
effect due to cineole only included 8% of species with growth reductions and 3%
with an increase. The camphor only effect alternatively had 25% of species with
significant growth reductions to leaf area and no significant increase. Both treatments
also demonstrated a significant effect in 28% of species showing significant
reductions in leaf area, 3% of species with a significant increase and 3% of species
with a significant growth reduction for cineole and an increase for camphor. No
significant effect on leaf area of the assemblage was found for 28% of species for leaf
area of the sampled assemblage.

108

(a).
92+
2 Ci-Ca+

1 02+

16

n = 36

14
differences

12

24+

10

8
6
4
2
0
'Cineole' o nly

'C am p hor' only

Both Trea tm en ts
Significant

No Significant
D ifference

Tr e atm e nt Significance to 'Contr ol'

Figure Xa. Total number of tree, shrub, vine and herb


species showing significant 'shoot length' variation in the
post-emergent measurements.

(b).
18
No. of species with significant
differences.

No. of species with significant

18

16

n = 36

14

62+

12
10
8

73+
1Ci-Ca+

13

31+

6
4
2
0
'Cineole' only

'Cam phor' only

Both Treatm ents


Significant

No Significant
Difference

Treatm ent Significance to 'Control'

Figure Xb. Total number of tree, shrub. vine and herb


species showing significant 'radical length' variation in the
post- emergent measurements.

109

(c).
16

No. of species with significant


differences

18
16

Note: analysis not available for one species.

n = 35

14
12
10

101+

42+

2-

6
4
2
0
'Cineole' only

'Camphor' only

Both Treatments
Significant

No Significant
Difference

Treatm ent Significance to 'Control'

FigureX. Total number of tree, shrub, vine and herb species


showing significant 'leaf number' variation in the post(d).emergent measurements. (note: analysis not available for one species)

No. of species with significant


differences

18
16
14

Note: analysis not available for one species.


n = 35

12
10
8

101+
1Ci-C a+
10

931+

6
4
2
0
'Cineole' only

'Camphor' only

Both Treatments
Significant

No Significant
Difference

Tr e atm e nt Significance to 'Contr ol'

Figure X. Total number of tree, shrub, vine and herb


Figure
19. Total number of tree, shrub, vine and herb species
species showing significant 'leaf area' variation in the postshowing
significant variation in the post-emergent measurements
emergent measurements. (note: analysis not available for one species)

compared to the control for shoot length (a); radicle length (b);
leaf number (c); leaf area (d).

For the trees and shrubs, shoot length (Figure 20a) was most affected with 83% of the
species having a significant response to the extracts. The significant effect on radicle
length, leaf number and leaf area for the trees and shrubs was less than that of shoot
length but displayed a similar high trend. This included 54% of species for radicle
length (Figure 20b), 56% for leaf number (Figure 20c) and 69% for leaf area (Figure
20d).

Where both treatments significantly influence growth at the same time, this was also
high with shoot length at 25% of species, radicle length at 33%, and leaf number and
leaf number at 30%. The response to the extracts for shoot length in the trees and
110

shrubs also included both significant reductions and increases in growth. Shoot length
had 46% of species with growth reductions, 33% with increases and 4% with a
reduction for cineole and an increase for camphor. For radicle length these growth
variations included 43% of species showing significant growth reductions and 13%
with significant growth increases. In growth variations for leaf number 43% of the
species had significant reductions and 13% had significant growth increases. For leaf
area 61% of species had leaf area reductions and 2% had leaf area increases.

No. of species with significant


differences

(a).
14

82+

12

4+

n = 24

32+
1 Ci- Ca+

10

6
4
2
0
'Cineole' only

'Camphor' only

Both Treatments
Significant

No Significant
Difference

Treatment Significance to 'Control'

Treatment
Significance
to Control
shrub species
showing
significant
Figure X. Number of tree and
'shoot length' variation in the post-emergent measurements.

No. of species with significant


differences

(b).
11

14
62+

12
10

n = 24

8
6

21+

2-

2
0
'Cineole' only

'Camphor' only

Both Treatments
Significant

No Significant
Difference

Treatment Significance to 'Control'

Figure X. Number of tree and shrub species showing significant


'radical length' variation in the post-emergent measurements.

111

No. of species with significant


differences

(c).
14

Note: analysis not available for one species.

12

n = 23

61+

10
8

10

22+

6
2-

4
2
0
'Cineole' only

'Camphor' only

Both Treatments
Significant

No Significant
Difference

Treatment Significance to 'Control'

Figure X. Number of tree and shrub species showing significant


'leaf number' variation in the post-emergent measurements.
(note: analysis not available for one species)

(d).

No of species with
significant differences

14
12
10
8
6

61+

n = 23

621+

4
2
0
'Cineole' only

'Camphor' only

Both Treatments
Significant

No Significant
Difference

Treatment Significance to 'Control'

Figure X. Number of tree and shrub species showing significant


Figure 20. Number of tree and shrub species showing significant
'leaf area' variation in the post-emergent measurements. (note:

variation
in theforpost-emergent
analysis
not available
one species)

measurements compared to the


control for shoot length (a); radicle length (b); leaf number
(c); leaf area (d).

All four growth parameters measured in the herbs (Figure 21a to d) were significantly
influenced by the extracts with 89% of the species for shoot length and 78% for radicle
length, leaf number and leaf area individually. The most significant response group for the
herbs was the effect where both extract treatments influenced growth of an individual
species for shoot length and leaf area. For radicle length and leaf number the effect of both
treatments was similar to the individual significant responses to cineole only and
camphor only, being lower in the number of species affected than for shoot length or
leaf area. These latter individual responses also showed some similarity in the number of
species influenced with the exception of camphor only for radicle length where the

112

(a).

No. of species with significant


differences

14
12

n = 9
51 Ci-Ca+

10
8
6
4

1-

1-

2
0
'Cineole' only

'Camphor' only

Both T reatments
No Significant
Significant
Difference
Tre atme n t Sig n ifican ce to 'Co n tro l'
species
showing
significant 'shoot
Figure X a. Number of herb
Treatment
Significance
to Control

length' variation in the post-emergent measurements.

No. of species with significant


differences

(b).

14
12

n = 9

10

31+

11 Ci-Ca+

6
4

1-

2
0
'C ineole' only

'C am phor' only

Both Treatm ents


S ignific ant

Tr e a tm e n t Sig n ific an c e to 'Co n tr o l'


Treatment Significance to Control

No Signific ant
D ifferenc e

F ig u re X b . Num ber of herb species showing significant 'radical


length' variation in the post-em ergent m easurem ents.

No. of species with significant


differences

(c).

14
12

n = 9

10
8
6
4
2
0

32-

2-

'Cineole' only

'Camphor' only

Both Treatments
Significant

Tre atme nt Significance to 'Control'

No Significant
Difference

Figure xb. Number of herb species showing significant 'leaf


number' variation in the post-emergent measurements.

113

differences

No. of species with significant

(d).

14
12
10
8
6
4
2
0

n = 9
31Ci-Ca+
2

21-

'C ineo le' only

'Camphor' on ly

Both T re atme nts


Significant
Tr e atm e n t Sig n ifica n ce to 'Co n tr o l'

No Significant
Difference

Treatment Significance to Control


Num
ber
herbspecies
speciesshowing
showingsignificant
significant 'leaf
F
ig
u
re
X
b
.
Figure 21. Number ofofherb
variation
area'
variation
in
the
post-em
ergent
m
easurements.
in the post-emergent measurements compared to the control' for

shoot length (a); radicle length (b); leaf number (c); leaf
area (d).

number of species was at its highest for that measured parameter. Although both
growth reductions and growth increases were recorded in all growth parameters the
majority of species demonstrated a reduction in growth i.e. 78% of species for shoot
length, 56% for radicle length, 78% for leaf number and 67% for leaf area.

The vines, Figure 22a to d, possessed the lowest number of species statistically
assessed with only 3 species represented. Significant effects of the extracts on growth
in the vines included 3 species for shoot length and radicle length individually, 1 for
leaf number and 2 for leaf area. Growth reductions accounted for all responses for
shoot length, with 1species for radicle length and leaf number individually and 2
species for leaf area. A positive increase in growth was only recorded in 2 species for
radicle length. Responses to cineole only, camphor only and the effect of both
treatments are represented across the 4 growth parameters measured, but due to low
sample size it was difficult to draw further information from the data.

For the assemblage as a whole the between treatment significance i.e. whether one
treatment has more effect than the other (Figure 23) demonstrated that the highest
proportion of species was influenced equally by both the cineole and camphor
treatments for radicle length, leaf number and leaf area. However, for shoot length the
difference between the treatments is clearly demonstrated where cineole has less
114

(a).

No. of species with

significant differences

14
12

n = 3

10
8
6

1-

11-

4
2
0

'C ineole' only

'C amphor' only

B oth Treatments
S ignificant

No S ignificant
D ifference

T re atme nt S ignificance to 'C ontrol'

Treatment
Significance
to Control
vine species
showing
significant 'shoot
Figu re X a. Num ber of
length' variation in the post-em ergent measurements.

(b).

No. of species with


significant differences

14
12

n = 3

10
8
6
1-

11+

1- 11+

2
0
'C ineole' only

'C amphor' only

Both Treatments
Significant

No Significant
Difference

T
re atme nt Significance
SignificancetotoControl
'C ontrol'
Treatment

Figure Xa. Number of vine species showing significant 'shoot


length' variation in the post-emergent measurements.

(c).

No. of species with


significant differences

14
12

n= 3

10
8

6
1-

4
2
0
'Cineole' only

'Cam phor' only

Both Treatm ents


Significant
Significant
T re atme nt Significance
to 'Control'

of vine species showing significant 'leaf


Figure Xc. NumberTreatment
Significance to Control
number' variation in the post-emergent measurements.

115

No Significant
Difference

Difference

No. of species with significant


differences

(d).
14
12

n=3

10
8
6

1-

1-

4
2
0
'Cineole' only

'Camphor' only

Both Treatments
Significant

No Significant
Difference

Treatment Significance to 'Control'

Treatment
Significance
to Control
of vine species
showing
significant 'leaf area'
Figure Xd. Number
Figure
22.
Number
of
vine
species
showing
variation in the post-emergent measurements. significant variation

in the
post-emergent measurements compared to the control for shoot
length (a); radicle length (b); leaf number (c);leaf area (d).

No. of species with significant


differences

25

21

21

19
16

20
14

'Cineole' > 'Camphor'

15

'Cineole' < 'Camphor'

7
7

10

'Cineole' = 'Camphor'

6
n n==35
35shoot,
shoot,radical
radiclelength
length
n n==34
34leaf
leafNo;
No;leaf
leafarea
area

0
s hoot length

radical length

leaf No.

leaf area

Significant Mean Variation Betw een Treatments

Total number
of tree,
shrub,
vine and
herbs
species
Figure
number
of tree,
shrub,
vine
and
herb showing
species showing
Figure
23.x.Total
significant variation between treatments for each growth parameter.
significant differences between treatments for each growth parameter.

effect than camphor and is nearly equivalent to the number of species equally
influenced by both extracts. In the case of cineole that significantly equals the
camphor effect in the sampled assemblage 46% of species were recorded for shoot
length, 54% for radicle length, 60% for leaf number and 62% for leaf area. For the
effect where cineole has less effect than camphor 40% were recorded for shoot
length, 26% for radicle length, 21% for leaf number and leaf area individually. In the
case where cineole has greater effect than camphor 14% of species were recorded
for shoot length, 20% for radicle length, 18% for leaf number and leaf area
individually.
116

For the trees and shrubs (Figure 24) a similar trend emerged for each of the growth
parameters statistically compared where the highest number of species responded
equally to the extracts, followed by cineole having less effect than camphor and
finally a small number of species where cineole had greater effect than camphor.
Proportionally, this equated to species where the significant effect of cineole and
camphor was equivalent occurring in 48% of species for shoot length, 52% for
radicle length and 59% for leaf number and leaf area. For the second highest level
where the cineole had a less significant effect than the camphor, 43% of species
were recorded for shoot length, 26% for radicle length and 23% for leaf number and
also leaf area. In the lowest response group where cineole had greater effect than
camphor 9% of species responded for shoot length, 22% for radicle length and 18%
for leaf number and also leaf area.

No of species with significant differences

14

13

13

12
12

11
'C in e o le ' > 'C a m p h o r'

10
10

'C in e o le ' < 'C a m p h o r'


'C in e o le ' = 'C a m p h o r'

A n a ly s is n o t a va ila b le

6
6

n =n 2=3 35
s h oshoot,
o t a n d radicle
r a d ic a l length
le n g th
n =n 2=2 34
le a fleaf
No .a
nd a
r e a area
No;
leaf

2
2

2
1

0
s h o o t le n g th

r a d ic a l le n g th

le a f No .

le a f a r e a

S ig n if ic a n t M e a n V a r ia t io n Be t w e e n T r e a t m e n t s

F ig u re X . N um b e r o f tre e a nd s hrub s p e c ie s s ho w ing s ig nific a nt va ria tio n b e tw e e n


tree
shrub
species
showing
Figure 24. Numbertreof
a tm
e ntsand
fo r e
a c h g ro
w th p a ra
m e te r. significant

differences between treatments for each growth parameter.

In the herbs (Figure 25), the largest number of species showed cineole had a
statistically equivalent effect to the camphor. The highest numbers were recorded
for radicle length, leaf number and leaf area while shoot length had a similar number
of species to the other treatment comparisons for that growth parameter. The
equivalence in treatment effect was 33% of species for shoot length and 56% for
radicle length, leaf number and leaf area. In the other response group comparisons
where cineole has a less significant effect than camphor and cineole has a greater
significant effect than camphor, the number of species responding was found to be
similar. For the response where cineole is less than camphor the proportion was
33% of species for shoot length and radicle length and 22% for leaf number and leaf
117

area. Where cineole has greater significant effect than camphor, 33% of species
were recorded for shoot length, 11% for radicle length and 22% for leaf number and
also leaf area.
14

'C in e o le ' > 'C a mp h or'


'C in e o le ' < 'C a mp h or'

10
differences

No. of species with significant

n = 9
12

'C in e o le ' = 'C a mp h or'

8
6
4

5
3

3
2

0
s hoot length

radic al length

leaf No.

leaf area

Sig n ifican t M e an V ar iat io n Be t w e e n T r e at m e n t s

F ig u re X . Num ber of herb species showing significant variation


Figure
25. Number of herb species showing significant
between treatm ents for each growth param eter.
differences between treatments for each growth parameter.

For the vines (Figure 26) where sample size was lowest i.e. 3 species, the highest
number of species were recorded where the significant effect of cineole equals that
of camphor. Further evaluation of this data was not attempted due to low sample

'C in e o le ' > 'C a m p h o r '

14
12
differences

No of species with significant

size as this identified that it was not representative of the vines per se.

n =3

'C in e o le ' < 'C a m p h o r '


'C in e o le ' = 'C a m p h o r '

10
8
6
4
2

0
s h o o t le n g t h

ra d i c a l
le n g t h

le a f N o .

le a f a r e a

S i g n if i c a n t M e a n V a r ia t io n B e t w e e n T r e a t m e n t s

F i g u r e X . N u m b e r o f vin e s p e c ie s s h o w i n g s ig n i fic a n t va ria t i o n b e t w e e n


of vine species showing significant differences
Figure
26. Number
t re a t m e n t s fo r e a c h g ro w t h p a ra m e t e r .
between treatments for each growth parameter.

The t-test comparisons (Appendix 11 and 12) also clearly demonstrate the extent of
the influence that the cineole and camphor treatments has on the individual species
when the number of significant effects impacting on shoot length, radicle length, leaf
number and leaf area are compared within and between species.

118

The numbers of significant responses of each species as regard the growth parameters
i.e. shoot length, radicle length, leaf number and leaf area have been grouped
irrespective of whether they were the same parameter or different parameters. This
was to provide an indication of how acute the influence was on the species. The
significant response to the cineole treatment included all 4 growth parameters for A.
caerulea, C. bartramia, *L. lucidum, L. confertus, and O. aemulus; 3 growth
parameters for C. hypoglauca, E. intermedia, E. saligna, G. semiglauca, H. flavum, L.
longifolia, M. tanarius, *P. clandestinum, S. capsicoides, S. luehmannii, S.
paniculatum and V. hederacea; 2 growth parameters for *A. cordifolia, *C.
coleutioides, C. parvifolia, F. coronata, O. populifolius, *P. dilatatum, and T.
australis; 1 growth parameter for A. melanoxylon, A. torulosa, *C. grandiflorum, *C.
camphora (from soil) and P. pupurescencs; and no growth parameters for A.
cunninghamii, C. enervis, H. bracteatum, and L. pulchella.

For the significant effect of the camphor treatment the 4 growth parameters were
influenced for C. hypoglauca, C. viminalis, C. enervis, F. coronata, L. longifolia, L.
confertus, S. paniculatum and V. hederacea; 3 growth parameters for *C.
grandiflorum, C. bartramia, E. saligna, *L. lucidum, O. aemulus, *P. clandestinum,
S. capsicoides, and T. australis; 2 growth parameters for

*C. coleutioides, H.

bracteatum, L. pulchella, O. populifolius and *P. dilatatum; 1 growth parameter for


A. caerulea, *A. cordifolia, C. parvifolia, H. flavum, E. intermedia, S. luehmannii and
P. pupurescencs; and no growth parameters for A. melanoxylon, A. torulosa, A.
cunninghamii, *C. camphora (from soil), G. semiglauca and M. tanarius.

The difference in the influence of the two extract treatments on growth is also
demonstrated, Appendix 11 and 12, with all 4 growth parameters varying significantly
between the cineole and camphor treatments for A. caerulea, F. coronata and *L.
lucidum; 3 parameters for C. enervis, E. intermedia, E. saligna, G. semiglauca, H.
flavum, L. pulchella, L. longifolia, *P. clandestinum, S. luehmannii and S.
paniculatum; 2 parameters for A. cunninghamii, *C. camphora (fresh from tree), C.
bartramia, O. aemulus, S. capsicoides and T. australis; 1 parameter for A. torulosa,
*C. grandiflorum, *C. coleutioides, *C. camphora (from soil), C. hypoglauca, C.
parvifolia, M. tanarius, L. confertus, O. populifolius, *P. dilatatum, V. hederacea and

119

P. pupurescencs; and no differences between treatments for A. melanoxylon, *A.


cordifolia and H. bracteatum.

2.5 Discussion
2.5.1 Vascular Plants
The germination response for the vascular plants (Figure 8) demonstrated much
variation in the percentage of seed germinating for each species. At least half the
species attained a germination percentage of 50% or higher, with a third showing high
germination exceeding 80%. Low germination percentages for some species were
attributed to seed dormancy, low seed viability or lack of time allowed for
germination completion at the end of the trial.

Viability was not tested with

tetrazolium as many of these seeds were extremely small or difficult to locate and
extract from the media.
In this glasshouse experiment the results of the deviance tests (log-likelihood ratio)
applied to statistical model produced from the Richards function for the asymptote
i.e. the total number of seeds germinated, mid-inflection point i.e. the number of
seeds where germination begins to slow, and germination rate of all vascular species
revealed no significant effect due to the cineole or camphor treatments. However,
the deviance test did demonstrate that a significant effect existed for the days to midinflection i.e. a delay to the time elapsed until germination began to slow. This
identified that the significant effect on vascular plant germination over time is that it
is delayed by the treatments. Further statistical assessment using multi-level analysis
identified that this effect was due to both the cineole and camphor extracts with no
significant difference between these treatments. This is consistent with the prestatistical analysis of the commencement time of vascular plant germination (Figure
9a) and completion time of germination (Figure 9b) where the delay is similar for
both cineole and camphor treatments. Comparison of the residual analysis of the
days to mid-inflection (Appendix 10, Figure A10.2) with the modeled data from the
Richards function (Appendix 8) revealed that the most influenced species in
ascending order of effect of the treatments for the time taken to germinate including
which treatments were of influence (Table 11).

120

These data indicate that camphor laurel allelopathy has three distinct effects on the
time taken for germination i.e. stimulation, delay or no effect, which is variable
depending on the species of vascular plant and to a much lesser extent the chemo-type

Table 11. The most influenced vascular species by the treatments in the time taken to
reach mid-inflection of germination.

Species

Effect of cineole

Effect of camphor treatment

treatment

M. tanarius

no influence

reduced

S. capsicoides

increased

increased

F. coronata

reduced

reduced

C. bartramia

increased

increased

G. cymosum

increased

no data

O. aemulus

reduced

reduced

*C. camphora (from soil)

increased

increased

*P. dilatatum

increased

increased

C. enervis

reduced

reduced

*L. lucidum

increased

increased

A. caerulea

reduced

reduced

M. philippensis

reduced

reduced

L. longifolia

increased

increased

C. hypoglauca

increased

increased

H. flavum

increased

increased

*A. cordifolia

increased

increased

of the camphor laurel tree. The effect on a plant assemblage, particularly where a
delay to germination over time occurs would be a reduction in plant competiveness as
faster germinating species would be advantaged. Ultimately, this would result in a
very different vegetation composition over time than if the native species only were
present. However, post-germination effects also need to be assessed as growth
following germination is also substantially altered.
121

For the growth of shoot length, radicle length, leaf number and leaf area (Appendix 11
and 12) 36 species of the original 54 species were tested by t-tests of the postemergent measurements. For all the vascular species tested i.e. the sampled
assemblage as a whole (Figure 19a to d) the t-tests demonstrated that the significant
responses to both the applied extracts were 86% of species for shoot length, 64% for
radicle length, 54% for leaf number and 71% for leaf area with the majority of species
displaying reductions in growth, identifying a highly significant proportion of species
and growth parameters influenced by the extracts. The trend was found to also be
relatively consistent in the life-form grouping of trees and shrubs, herbs and vines
(Figure 20a to d; 21a to d; and 22a to d). This high but differential response by species
where growth is influenced in both a positive but mainly negative fashion, where
species may or may not be influenced by allelopathy and where various parts of the
plant are influenced to a lesser or greater extent depending on the species is consistent
with the findings of Asplund (1969), Firth (1979) and Chou et al. (1989). In the study
undertaken by Firth (1979) where aqueous extracts from an unknown chemo-type of
*C. camphora were applied to the seeds of lambs tongue (*Plantago lanceolata),
Wimmera ryegrass (*Lolium rigidum) and catsear (*Hypochoeris radicata) reductions
in germination percentage occurred but without any effect on seedling vigour, shoot
length, leaf number or colour. Considering that this previous experiment was only
continued for a two week period compared with months in some cases for this current
experiment, it has been shown that the time exposure to the extracts is important in
demonstrating the overall influence of the effect. Johns (1994) further investigated the
effects of *C. camphora extract of unknown chemo-type on the germination of kikuyu
(*P. clandestinum), *Cedrela odorata, Toona ciliata var. australis, Casuarina
cunninghamiana, Euodia elleryana as well as *C. camphora allowing most seed to
fully germinate for up to two months. The study found that the extracts of *C.
camphora did not significantly stimulate or inhibit the germination of its own seed but
did inhibit germination of the other species tested, excluding Euodia elleryana which
failed to germinate. In this current experiment the fresh seed of *C. camphora was
subjected to extract treatment but could not be used for analysis due to a desiccation
event in the control. This could not be repeated due to an absence of fresh fruit on
the trees and older seed originating from A soil horizon had to be used. However, the
germination profile for this older soil exposed seed showed a marked decrease for
both the cineole and camphor extracts applied (Appendix 1, Figure A1.17) with
122

the camphor possessing the lowest germination. The deviance and multi-level
analysis showed that a significant effect occurred only with the days to midinflection point of germination i.e. delaying germination with no significant
difference between the cineole and camphor treatments. The test identified that the
total number of seeds germinating, germination rate and the number of seeds
germinating at the inflection point of germination remained unaltered. The t-tests
revealed that the cineole extract significantly reduced radicle growth of the seedlings
of its own species below the control. A significant difference was also found to exist
between the cineole and camphor treatments with cineole having a greater
suppressive effect than camphor. Although this does not provide evidence of the
way fresh seed of *C. camphora behaves when exposed to extracts it does identify
that a suppressive effect does occur for older seed within the soil profile.

In the t-test comparison for all vascular plants to determine whether cineole or
camphor had greater effect, the highest numbers of species were influenced equally
by both treatments (Figure 19a and b). However, where the cineole effect is greater
than camphor or where it is less than camphor these responses are also
represented, but are on average less than half the number of species influenced by
both treatments. The only variation on this effect is shoot length where the effect of
cineole has less influence than camphor. The variation is nearly equivalent to the
number of species influenced equally by both treatments. Examination of the
individual life-form groupings (Figures 13a to d, 14a to d, and 15a to d) also supports
that variation occurs across the life-forms due to differential effect. The effect of the
treatments indicates that most vascular plants are influenced by both extracts but with
shoot length being influenced more by camphor than cineole. What this also
means is that the process of alpha selection in the local plant population which is
invaded by camphor laurel not only involves a delay to the time taken for
germination, but also a reduction in growth following germination. For some less
robust species this process may result in a severe reduction in abundance over time.

When the post-emergent t-test groupings for growth are compared with seed
characteristics (Table 6), allelopathy was not found to be influenced by the physical
attributes of the receptor seed such as size, moisture content of the pericarp, seed

123

dormancy, life-form or plant family. This may suggest that the allelopathy operates in
an innate fashion possibly relating to the chemistry of the seed but this is unknown.

Of the 54 vascular species tested in the experiment (Table 6) 46 species were found to
be suitable for the statistical analysis of germination over time. The species excluded
from the analysis of germination due to errors in methodology or poor germination
response included C. rubra, *L. camara, *P. suberosa, A. excelsa, J. pseudorhus, F.
xanthoxyla, D. australis, D. pruriens and C. australe. For *C. camphora (fresh from
tree) germination did not occur in the control and zero values were recorded making
statistical analysis impossible for comparisons of the treatments to the control. The
absence of germination in the control was most likely due to a desiccation event
during germination as some drying of the media was observed during the germination
process. Further information that supports this is the experiment by Johns (1994)
where camphor laurel extract was not found to significantly influence the germination
of its own fresh seed. Furthermore, the work of the U.S. Dept of Agriculture (1897),
Troop (1921), Ghosh (1977) and Panetta (2001) identified that fresh camphor laurel
seed has a high viability without any extract application, and Chen et al. (2004) found
that this high viability decreases as moisture stress occurs. For some species statistical
comparison was not possible in the t-tests due to zero germination, missing data or
damage to seedling due to rodents. In the case of A. littoralis phyllode number and
phyllode area were not possible to calculate due to the absence of these parameters for
measurement. Predation of germinating seed by rodents occurred in the cineole
treatment of C. viminalis limiting statistical comparison to only the effect of the
camphor treatment.

2.5.2 Soil Algae


In this algal glasshouse experiment the results of the deviance tests (loglikelihood
ratio) applied to statistical model produced from the Richards function for the
asymptote i.e. growth coverage of the media, mid-inflection point i.e. the point
where growth begins to slow, and growth rate of the algae revealed no significant
effect due to the cineole or camphor treatments. However, the deviance test did
demonstrate that a significant effect existed for the days to mid-inflection i.e. a delay
to the time elapsed till growth began to slow. This identified that the significant effect
on algal growth over time is that it is delayed by the treatments. Further statistical
124

assessment using multi-level analysis identified that this effect was similar for both
the cineole and camphor extracts with no significant difference between these
treatments. Comparison of the residual analysis of the days to mid-inflection with
the modeled data from the Richards function (Appendix 10), revealed that the algae
cultures were influenced by the treatments for the time taken to grow over the media
including which treatments were of influence. This included the 3 algae cultures
trialed, all of which had an increased period for coverage of the media by both the
cineole and camphor treatments.

This delay to the coverage of the punnets by soil algae was most profound and is
clearly demonstrated in Figure 12a to c. where three separate algal cultures were
propagated and grown. Both the cineole and camphor treatments resulted in a
significant time delay to the coverage of the media but with no significant difference
between the influences of the two treatments as assessed by the deviance test and
multi-level analysis. This algal experiment involved 20 species of cyanobacteria
(Table 10), many of which were found to be either delayed in growth, severely
reduced in number or absent from the media in which the camphor laurel extracts
were applied. Diversity of algal species at the time of full media coverage
demonstrated that the number of species was reduced to 39% for the cineole
treatment and 54% for the camphor treatment compared to the control. This loss of
diversity is also clearly shown in Figure 14 to 18, where many species are either
severely

decreased

in

abundance

such

as

Cylindrocapsa,

Closteridium,

Pseudotetraspora, Pamellopsis, Nodularia and Pseudanabaena; or absent such as


Ulothrix, Oedocladium sp.1, Oedocladium sp.2, Haematococcus, Entophysalis sp.1,
Gloechaete, unnamed Cyanobacteria sp.21, Microchaete and Cylindrospermum. Only
five algal species increased in abundance such as Calothrix, Spirulina, Phormidium,
Amphora and Anabaena.

The loss of algal diversity in the camphor and cineole' treatments is of note,
especially in light of the fact that soil algae are important for maintaining soil
function. The presence of soil algae increases wetability and water retention capacity
(Booth 1941; Whitton and Potts 2000; Buscot 2005; Issa et al. 2007), mineralization
and humification processes (Whitton and Potts 2000; Buscot 2005) such as carbon
and nitrogen fixation (Shields and Durrell 1964; Mayland and McIntosh 1966; Rice
125

1992, Lange et al. 1994; Whitton and Potts 2000; Philippot and Germon 2005; Issa et
al. 2007). In some situations algae also increase iron, sulphur, phosphorus and
calcium availability (Whitton and Potts 2000), soil structural aggregation (Buscot
2005; Issa et al. 2007), soil erosion prevention (Booth 1941; Shields and Durrell
1964), plant succession enhancement (Shields and Durrell 1964) and as a food source
for invertebrates (Edwards 2000; Maraun et al. 2003; Buscot 2005). In work by Roger
and Burns (1994) significant increases to seedling emergence was found when soils
were inoculated with cyanobacteria. Inoculation has also been found to encourage soil
microbial population and increase organic material, nutrient availability and soil
stability in studies by Ashley and Rushforth (1984) and Johansen et al. (1994). These
ecological soil functions provided by algae suggest that the effect to soil structure,
moisture, nutrient availability, invertebrate food supply, and seedling response may be
substantially altered through allelopathic influences to soil algal composition and
growth over time from decaying camphor laurel leaves. It indicates that the reduction
of algal diversity and vigor may have an indirect allelopathic effect on seedling
germination and growth below the camphor canopy through possibly altering some
soil processes such as moisture retention and nutrient availability. Although soil fungi
were not investigated during this study previous research on the fungicidal effects of
*C. camphora and related species by Abivardi (1979), Tiwari and Dixit (1994), Chao
et al. (2000), Rahman (2000), Liu et al. (2001), Ali et al. (2002) and Ranasinghe et al.
(2002) found that the essential oils from this plant genus do sterilize or alter soil
fungal composition. Soil bacteria are also influenced by the allelochemicals from *C.
camphora as demonstrated in a study by Choi et al. (1999). Therefore, it is highly
likely that the soil fungi and bacteria in northern New South Wales are also affected
by the allelopathic extracts of camphor laurel. This would further add to any
influences to soil processes resulting from the algae being affected.

The deviance and multi-level analysis showed that a significant effect occurred only
with the days to mid-inflection point of algal growth i.e. delaying growth with no
significant difference between the cineole and camphor treatments. The test
identified that the coverage of the media, growth rate and algal coverage at the
inflection point of growth remained unaltered. Importantly, it also indicates that
species diversity of soil algae may be strongly influenced.

126

2.5.3 Environmental Control and Experimental Limitations


Environmental variables for this experiment were rigorously monitored and controlled
in an effort to minimize nutrient, temperature, moisture and allelochemical
fluctuations. The nitrogen and phosphorus adjusted extract solutions for the cineole
and camphor treatments (Figure 6a and b) demonstrated a close nutrient balance for
the vat solutions prepared. This included the use of four separate vat solutions to
ensure that allelochemicals did not volatilize or alter substantially. An initial error in
nutrient adjustment was rectified and that solution plus the treated seeds were
discarded. The pH of the solutions was adjusted at regular intervals due to a change in
the chemistry of the liquids over time (Figure 7a to d) providing a relatively stable
pH. The propagation unit (Figure 4) provided for adequate protection of the
treatments through enclosure from extract washout from glasshouse misting,
confinement of volatile loss as would be anticipated in a deep leaf litter and the
reduction of desiccation from high temperatures. The recorded temperatures (Figure
5) during the experiment revealed 6 high temperature peaks of 45C to 47C in
summer being brief due to shade cloth installation at these critical seed germination
temperatures. These hot periods lasted a few days at a time during the 360 days of the
glasshouse trial and any effect on germination was consistent across all species and
treatments. Hartmann and Kester (1968) indentified that seed from different species
can vary in the ability to withstand high or low temperatures with each having a
minimum, maximum, and optimum for germination. When temperatures exceed the
optimum range somewhere between 30C to 45C depending on species, germination
rate decreases and heat injury occurs if shade is not provided. In this trial
germination rates were identified as being high for many species or at least
statistically comparable for greater than 85% of species indicating that the high
temperatures did not adversely influence germination. Desiccation and heat scorching
was kept to minimal levels in the propagation units through:

the enclosure of the propagation area with shade cloth;

increased humidity due to enclosure of


polyshade,

127

propagation units with

regular application of extracts to the punnets; and

mist drift onto the top of the propagation units.

However, some punnets such as the control treatment of *C. camphora (fresh from
tree) at the early stages of the glasshouse trial were prone to desiccation resulting in
poor germination. It was at this stage that enclosure of the punnets in moisture
confining propagation units was applied. Minimum temperatures reached 5C for a
brief period in mid-winter resulting in a slowing of germination and growth in the
punnets with no adverse effects noted.

Some limitations in the glasshouse research have been identified as:


an inability to generate standard errors for the r2 values of the Richards
function for the non-linear regressions performed on the germination
and algal growth data. Brooks (pers. com. 2010) suggested that this
would have only been required if the phytochemical effect was
assessed for an individual species not the group as a whole as in this
case;
small replication size due to the large number of seeds required for the
treatments and limited availability of seed due to seasonal fruiting;
lack of repeatability in the glasshouse trial due to time constraints of
the research;
lack of glasshouse environment control resulting in high temperatures
although ameliorated by misting, shading and humidity control;
possible chemical differences created by aqueous extract preparation
for the glasshouse trial compared to leaf leaching and volatilization in
the field situation. The approach taken by Leicach et al. (2009) for
bioassays, particularly where volatile compounds such as cineole are
involved requires consideration as many do not dissolve in water and
only become allelopathic when bound to soil particles through aerial
transfer; and

128

constraints to allelopathy trials using germination. Leicach et al. (2009)


identified that germination bioassays although easier to perform, do
have limitations which include variation in sensitivity of germinating
seed to allelochemicals depending on species, while others are more
sensitive in the seedling stage or later growth.
Future experimentation requires consideration of these possible limiting factors to
improve ongoing research.

2.6

Conclusion

The glasshouse experiment demonstrated that a significantly high number of common


native plants and woody weeds are influenced by allelopathic compounds originating
from *C. camphora leaves. These may include a range of monoterpenes, essential oils
and possibly phenols. Phytoxicity was found to significantly influence the time taken
for vascular plants to germinate with no significant differences between the cineole
and camphor chemo-types. No significant effect was found for the total number of
seeds germinating, the number of seeds germinating at the point where germination
begins to slow and the germination rate. The allelopathy produced early or late
germination responses without any influence resulting from seed attributes such as
size, moisture content of the pericarp, dormancy, life-form or plant family.

The reduction in time taken to germinate was further identified to significantly


influence the growth of most seedlings with both reductions and stimulations to shoot
length, radicle length, leaf number and leaf area. However, reductions in growth were
most commonly found for the species tested, indicating that extracts from the leaves
of *C. camphora have the potential to mostly suppress growth of the surrounding
plant assemblage while at the same time stimulate or have no effect on some species.
The proportion of effect attributed to either the cineole or the camphor treatments
was found to be relatively equivalent for the majority of species tested. Less than half
the number of species demonstrated variation to this where the cineole effect is
either greater than camphor or where it is less than camphor. The similarity of
effect showed consistency for radicle length, leaf number and leaf area not only for
the assemblage as a whole but also for the life-form groupings. The only variation to
129

this was shoot length where the effect of the camphor treatment exceeded the
cineole treatment being nearly equivalent to the effect where cineole is equal to
camphor. The variations in effect suggest that although the influence of the two
chemo-types on seedling growth is relatively equivalent for most growth parameters,
shoot length is influenced to a greater extent by the camphor extracts than cineole.

Ultimately, the findings indicate that direct allelopathy may be in operation below the
camphor laurel canopy where seed germination and seedling growth are influenced
directly by the allelochemicals with little differences between the two chemo-types.
This two stage alpha selection is possibly exacerbated by the trees dense canopy and
shallow, highly competitive root structure which would place further competitive
stress on seedlings affected by allelopathy and possibly alter the successional
sequence of plants regenerating below the camphor laurel canopy, particularly for less
robust species. Verification of these allelopathic influences to germination and growth
from the glasshouse trial are now required under field conditions to fully identify if
this effect is substantial enough to change succession in the local plant communities.

The glasshouse experiment demonstrated that soil algae are influenced by allelopathic
compounds originating from *C. camphora leaves. These may include a range of
monoterpenes, essential oils and possibly phenols. Phytotoxicity was found to
significantly influence the time taken for the growth of soil algae, with no significant
differences between the cineole and camphor chemo-types. No significant effect was
found for the coverage of the media, media coverage at the point where growth begins
to slow and the growth rate. Soil algal diversity however, did respond with dramatic
reductions in the cyanobacteria indicating that soil function such as moisture, nutrient
and humification processes could be influenced. The literature also indicated that the
abundance and diversity of soil fungi, bacteria and invertebrates may also be altered
by this allelopathy, further adding to the possible negative impacts to soil function.

Reductions in growth were found for the algal species tested, indicating that extracts
from the leaves of *C. camphora have the potential mostly to suppress growth of the
surrounding algal assemblage while at the same time stimulate or have no effect on
some species. The proportion of effect attributed to either the cineole or the
camphor treatments was found to be similar for the majority of species tested.
130

Ultimately, the algal findings suggest that an indirect allelopathy may be in operation
below the camphor laurel canopy where soil microbial populations and soil function
are influenced by camphor laurel extracts with possible influence on vascular plant
seed germination, with little differences between the two chemo-types.

In summary the findings that address the hypothesis indentify that:

there is significant effect on the germination and growth of vascular plants and
soil algae from the regeneration assemblages under glasshouse conditions
resulting from camphor laurel leaf extracts, and

there is no significant difference in allelopathic effect due to the chemo-types


on seed germination, early plant growth and algal populations from the
assemblages.

Two studies are now required before these findings can be verified as active in the
field situation. Firstly, a technique of field chemo-type differentiation is needed which
enables an inexpensive, rapid and reliable identification without the aid of expensive
and time consuming gas chromatography. This would enable a large number of
camphor laurel trees to be surveyed and statistical validity provided through a large
sample size. Secondly, a field assessment of seedlings comparing growth below the
camphor laurel canopy of both chemo-types using the developed identification
technique and other vegetation to indentify if allelopathy is active in the field, and
whether the glasshouse findings are relevant.

131

-Chapter 3:Field Recognition of the Camphor Laurel Chemo-types.

3.1 Introduction
To undertake the field comparison of germination and growth of regeneration
assemblage species under the cineole and camphor chemo-types identified by
Hirota and Hiroi (1967), Stubbs and Brushett (2001) and Stubbs et al. (2004), a
considerable number of camphor laurel trees must be sampled to establish
repeatability and statistical validity of the field sampling for this study, as well as the
ultimate comparison to the allelopathy hypothesis generated from the glasshouse trial.

The work by Hirota and Hiroi (1967) used olfactory recognition of volatiles liberated
from the leaf, while Stubbs and Brushett (2001) and Stubbs et al. (2004) applied gas
chromatographic (GC) analysis to provide highly accurate assessments of individual
oils of the leaf, bark and wood.

Both these techniques possess application problems for broad-scale field assessments
of the chemo-type, particularly ecological studies where large numbers of trees need
be sampled for statistical purposes. The technique of aroma interpretation is based on
a well developed sense of smell of the individual chemo-type, difficult for the novice
leaf smeller and requiring a lengthy period of olfactory acclimatization to enable
accurate interpretation of the volatiles.

This aroma interpretation also has the added problem of not being able to be
adequately demonstrated without a standard comparative smell to which users could
refer, and although not impossible to replicate would create problems of practical
repeatability without a GC standard for comparison.

The technique of gas

chromatography on the other hand requires relatively time-consuming laboratory


procedures and specialized skills in the use of equipment and interpretation of the
data. As a result, gas chromatography would become expensive and cumbersome
where large numbers of trees are to be differentiated for their chemo-type, especially

132

across numerous sites. A technique of chemo-type differentiation was required which


not only possessed consistency and repeatability, but is inexpensive, applicable in the
field, can be applied at any time of year, is not time consuming, and require minimal
specialized equipment and analytical skills.

3.2 Aim
The aim of this chapter is to identify any internal or external morphological, secretory,
chemical or olfactory characteristics of the camphor laurel leaf which could be used to
differentiate the cineole and camphor chemo-types reliably, inexpensively and
easily in the field.

3.3 Methods
The methods involved five separate analytical tests, each with an initial pilot trial, to
establish a worthwhile research direction. These comprised:

sectioning tests of the lamina and application of various histological


stains to identify possible microscopic differences in chemical
histology or morphology between the chemo-types;

leaf chlorophyll extraction, solution colour comparison of the plastid


pigments and precipitation of the separated residues;

study of the venation of the leaves from the two chemo-types;

study of leaf sclerophylly of the two chemo-types; and

olfactory recognition of the chemo-types using volatile leaf aroma


verified by gas chromatography (GC).
3.3.1 Sectioning and Staining

Fresh leaf material was gathered on a weekly basis for ongoing sectioning, staining
and microscopic examination. All leaves were placed in hermetically sealed plastic

133

bags to reduce transpiration in the field, and stored at 15C in a refrigerator following
collection.

A method of hand sectioning was adopted, applicable in the field situation as it does
not require the use of a bulky microtome, tissue embedding or freezing techniques.
Histologically stained leaf sections of the two chemo-types of *C. camphora were
compared at three locations on the leaf. These were: a transverse-section (TS) at midpoint of the leaf lamina, a TS and longitudinal section (LS) at mid-point of the petiole,
and the adaxial and abaxial phylloplane at mid-point of the lamina. Epidermal
structures of the leaf were prepared by cutting thin 10x5mm sections of the adaxial
and abaxial surfaces, as much of the external morphology was observed to be
microscopically visible without epidermal peels or leaf chlorophyll clearing methods.

In the sectioning of TSs of the lamina, an initial cut was made across the mid-point of
the leaf using a sharp blade. Single TSs using a single hand held blade were found to
be inadequate for the purpose of examining the oil globules of the secretory cells as
these occur infrequently and repetition was required. For this purpose, a twin blade
razor was employed to shave up to twenty TSs rapidly. This rapid methodology
provided long thin sections without the aid of a microtome.

Sectioning tests of leaf tissue using many histological stains were then undertaken at
three defined levels. The first involved the use of the cineole and camphor chemotype of camphor laurel which had previously been assessed by Stubbs and Brushett
(2001). Ten mature leaves were plucked randomly from the lower canopy of each
chemo-type for this test as mature leaves provided a well developed cellular structure
and chemical composition. Various histological stains, (Table 12) were applied in a
systematic procedure to establish if any visual differences existed between the two
chemo-types through variation in the staining of specific targeted cellular structures.
Sectioned leaf material was placed onto microscope slides and a few drops of the stain
applied to adequately cover the sections. A period of up to 30 minutes was allowed
before stain clearing or microscopic examination to allow adequate stain infiltration of
the tissues and, where applicable, excess stain was cleared with a solution of 70%
ethanol. Comparative examination under low and high power of the light microscope

134

including photo-microscopy was then performed for the internal and external leaf
morphology using the eleven identified stains. The measurement of cells or cellular

Table 12. Histological stains, substances stained and references to staining techniques used in
the systematic sectioning of camphor laurel leaves.
stain

substances stained

technique reference

Aniline Blue

callose

Ruzin (1999), University of Illinois (2006)

Toluidine Blue

lignin, tannins
polyphenols i.e.
pectin, pectic
substances.

Ruzin (1999), University of Illinois (2006)

Phloroglucinol

lignin

University of Illinois (2006), Labs-Mart (2006)

IK Iodine

starch, cellulose

Ruzin (1999), University of Illinois (2006)

Methylene Blue

cellulose

Ruzin (1999), University of Illinois (2006),

Ferric Sulphate

tannins

University of Illinois (2006)

Safranin Red

Chamberlain (1925), Ruzin (1999),University


of Illinois (2006), University of Berkeley (2006)

Sudan III

cutin, chromatin,
lignin, phenolics,
tannins, suberin
chloroplasts,
chitin, nuclei &
chromosomes.
oils, fats, cutin

Santos (1930), Labs-Mart (2006)

Red Henna

camphor?

Cartwright-Jones (2006), Chemindustry.com (2006)

glycogen

Chamberlain (1925), Ruzin (1999)

calcium oxalate

Pizzolata (1964)

(lawsone methyl ether)

Cochineal
(simple carmine)

Pizzolato

structures in microns (m) was further undertaken where visual differences appeared
worthy of further validation.

The second analysis included a blind test to establish if consistency in cellular


structure could be attained using the stain and targeted tissues identified from the first
test on four other gas chromatographed trees of the two chemo-types.

For the final test, staining of leaf sections from an additional ten trees and verification
by GC was undertaken to demonstrate the possible application of the technique to the

135

field population. GC work was undertaken on the trees at the Centre for
Phytochemistry at SCU using a Hewlett Packard HP6890 series GC system. The GCMS FID Bretts technique was applied with the advantage of being able to assess all
compounds present in the leaf. A standard column was used with the split ratio set at
25:1 and oven temperature start at 50C being ramped at 8C per minute until 300C
was achieved.

For the second and final tests, only three leaves were used from each tree to reduce
time taken for these assessments. Comparative cellular structures were then measured
in microns using the developed stain. The chemo-type identity of these trees remained
blind until the procedure of identification with the stain was completed. The
developed technique of sectioning, staining, photo-microscopic examination and
measurement of cells using the successful stain were then applied to this group of
trees. Following this second assessment and comparison, leaf samples from 10 other
trees were collected, assessed with the stain, then independently gas chromatographed
enabling a broader sampling and application of the stain to the camphor laurel field
population. This was to establish whether the differences identified using the first two
trees could be applied generally to other camphor laurel trees through graphical
comparison of the targeted cellular structures.

3.3.2 Leaf Chlorophyll Extraction


This pilot test involved the comparison of plastid pigments and solution precipitates
from the two chemo-types using a technique outlined by Howarth and Warne (1952)
for chlorophyll clearing from solution. Ten grams of finely cut mature leaves from
each of the two chemo-types were placed in 10mL of acetone to release chlorophyll,
volatile oils and plastid pigments to solution. To this filtered solution were added
10mL of petroleum ether and a few drops of distilled water to separate chlorophyll
from the plastid pigments and other compounds leached from the leaf by the acetone.
The chlorophyll-free solution from each chemo-type was compared with each other
for colour differences, further stained with a few drops of Sudan III, and separately
with Red Henna stain. Precipitates were also prepared using the unstained
components of the resultant chlorophyll solution and clearer plastid pigment solution
using watch-glasses to establish any differences in the precipitated residue such as

136

colour, gum or crystalline development. The test was repeated twenty times to
establish if differences existed between the two pilot trees.

3.3.3 Venation
For the pilot test using venation, ten mature leaves were examined from each gas
chromatographed chemo-type using back lighting and microscopic comparison to
determine if leaf clearing using nitric acid or other solvents would be applicable.
3.3.4 Leaf Sclerophylly
During the process of leaf gathering and sectioning it was noticed that differences in
leaf hardness and rigidity occurred in some trees. A pilot test was conducted to assess
whether sclerophylly could be applied to determine chemo-type. Eighteen GCd trees
were used which included seven cineole and eleven camphor chemo-typed plants.
A blind test for hardness and rigidity using the manipulation of the leaf between
thumb and fore-finger was investigated using the description of the leaf as either soft
(pliable with thin margins) or hard (leathery with thick margins). Success rate was
evaluated as a percentage during this pilot.

Further testing using micrometer measurements of the midrib to three decimal places
e.g. 0.001mm, were performed at mid-point of the lamina for each chemo-type. Ten
leaves were gathered from each of five gas chromatographed cineole trees and nine
camphor trees for this comparison. A single factor analysis of variance (ANOVA)
was applied to establish significance at the 0.05 level.
3.3.5 Olfactory Recognition
The olfactory test involved four short pilot studies to establish a possible technique
applicable to field recognition using the characteristic smell liberated by each chemotype. Five mature leaves were taken from two trees, one each of the camphor and
cineole chemo-types. These four pilot tests included crushing between the fingers
and smelling; cutting, placement and smelling in zip-seal plastic bags to enclose and
concentrate the volatiles; cutting and smelling using an aspirator with attached plastic
funnel and nasal mask; and cutting and placement in a wide porcelain dish with a
volatile trapping glass cover. Aroma was then compared to both the cineole and
camphor chemo-types of camphor laurel which had previously been gas

137

chromatographed in this study and by Stubbs and Brushett (2001) and Stubbs et al.
(2004).

Following the recognition of the most promising olfactory technique from the pilot
study, a blind test was applied to five mature leaves gathered from each of thirteen
trees of the camphor chemo-type and five trees of the cineole chemo-type which
had previously undergone GC analysis. This was repeated and an accuracy of the
technique was calculated as a percentage of successful olfactory recognition from one
hundred and eighty-one separate blind tests.

3.4 Results
3.4.1 Sectioning of Lamina
The substances stained and observed within the leaf structure of the camphor laurel
chemo-types using the various stains included callose, calcium oxalate, lignin,
tannins, starch, cellulose, cutin, glycogen, polyphenols and essential oils. This enabled
visual comparison of various cellular structures of the:

petiole, such as the epidermis, collenchyma, parenchyma, the stele


(xylem and phloem), stellar fibre bundles, laticifers (resin ducts);

external lamina, such as trichomes, stomata, epidermal cells, oil


secretory cells, mucilage secretory cells; and

internal lamina, such as palisade and spongy mesophyll, chloroplasts,


starch grains, vascular structures (xylem, phloem and associated
sclerenchyma fibre bundles), parenchyma containing calcium oxalate
(rhaphides), secretory cells and oil globules.

Of the various histological stains used, only one demonstrated possible differences
between the two chemo-types. This was Sudan III which stains essential oils, fats and
cutin red (Santos 1930, Labs-Mart 2006). The structures which emerged with the use
of this stain were the essential oil globules found only occasionally within the
secretory cells of both chemo-types and more commonly throughout the mesophyll

138

particularly immediately near the abaxial and adaxial epidermis. These structures are
compared in TSs of the lamina for the first two test trees, (Figure 27a and b). The
cineole chemo-type was initially found to possess very few large oil globules and
rhaphides while the camphor chemo-type had many small oil globules which
appeared to occur more frequently in the secretory cells. This was also demonstrated
in Table 13 where oil globule size was shown to be much reduced in the camphor
chemo-type in the first two trees studied. Further oil globule measurement in four
other trees (Table 14), and verification through blind testing against GC analysis, also
provided additional support for this variation being related to chemo-type. In the
broader application of the analysis to nineteen other trees including an additional ten
trees being gas chromatographed, three globule types emerged, i.e. those possessing
many small globules as in the initial camphor globule type, those possessing many
large globules as in the initial cineole type and those possessing both large and small
also of the camphor type (Figure 28). In the application of the technique to three
wild trees in a forest situation (Figure 29), the cineole chemo-type was identified as
being able to possess both large and small globules, placing no consistency on oil
globule size being related to chemo-type. The shape and colour of the oil globule was
found to vary slightly between the chemo-types in the initial sectioning. This was later
identified as not being consistent as the sample size in the study increased.

The analysis of chemo-type using gas chromatography of camphor laurel leaves of


both the camphor and cineole chemo-type revealed 19 other oils not recorded
previously.

These

included

-terpinene,

-terpinolene,

-terpineol,

ledene,

viridiflorol, -phellandrene, phellandrene, borneol, -sabinene, p-cymene, -pinene,


mentha 1,8-dien, -humulene, -cubebene, -selinene, -selinene, trans-sabinene
hydrate, cis-sabinine hydrate and trans-caryophyllene.

3.4.2 Leaf Chlorophyll Extraction


The stained and unstained plastid solutions, and the precipitates of the unstained
chlorophyll and plastids solutions of the two chemo-types were found to possess no
visual differences in colour, gum residue or crystalline development.

139

palisade mesophyll
cutinized layer
adaxial epidermis
- oil enriched, some oil
globules
oil secretory cell
large oil globule
- containing calcium
oxalate crystal
rhaphid cell
- containing prismatic
calcium oxalate
crystals
small oil globules
in the mesophyll
spongy mesophyll
air bubble artefacts
abaxial epidermis

- oil enriched, some oil

(a.) Cineole chemo-type

globules
cutinized layer

air space

air bubble artefacts


cutinized layer
adaxial epidermis
- limited oil enrichment
palisade mesophyll
- containing chloroplasts
chloroplast
small oil globules
oil secretory cell
spongy mesophyll
- containing chloroplasts
small oil globule
in the mesophyll
air space
abaxial epidermis
- limited oil enrichment
cutinized layer

(b.) Camphor chemo-type


Figure 27. Hand TSs of the (a) cineole and (b) camphor chemo-types of camphor
laurel stained with Sudan III showing possible oil globule variation later identified as
not being related to chemo-type.

140

Table 13. Comparison of secretory cells and oil globules of the leaf in two gas
chromatographed chemo-types of camphor laurel using Sudan III stain.
Tree 1 Cineole Chemo-type
Secretory cell
size (m)
Section No.

Length

Oil globule
size (m)

Diameter

Length

Secretory cell
size (m)

Diameter

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15

60
36
60
30
42
48
40
30
50
48
42
30
46
43
46

30
30
40
36
36
42
45
30
40
36
36
30
40
40
48

18
36
36
30
36
48
20
30
29
48
42
27
26
23
40

18
30
30
24
30
24
15
30
23
36
27
24
22
20
26

16

42

33

38

30

43.3

37

32.9

mean

Tree 2 Camphor Chemo-type

Length

25.6

Oil globule
size (m)

Diameter

Length

45
40
40
40
45
40
40
50
40
40
40
40
60

30
40
35
35
40
35
35
45
35
40
40
35
35

8
15
4
8
12
12
15
12
12
15
3
2
20

43

37

10.6

Diameter

8
15
4
8
12
12
15
12
12
12
3
2
20

10.4

Table 14. Comparison of the oil globules in four trees of camphor laurel whose
chemo-types were assessed using Sudan III staining, followed by verification through
blind testing against gas chromatography.
Oil globule size (m)
Tree 1
Section No.

1
2
3
4
5
6
7
8
9
10
mean
Comparison
to
Table 12.
Gas
Chromatography

Length

Diameter Length

13
18
15
16
15
18
20
18
18
15
16.6

13
12
15
10
12
12
10
12
12
15
12.3

Tree 2
Diameter

30
18
20
48
30
30
25

28.7

Tree3
Length Diameter

Tree 4
Length

Diameter

24
12
20
37
30
24
20

25
12.5
37.5
37.5
30
50

25
10
32.5
27.5
25
37.5

15
10
15
7.5
12.5
15
12.5
3.75
17.5
20

12.5
7.5
15
7.5
10
12.5
12.5
3.75
12.5
20

23.9

32.1

27.9

12.9

11.4

camphor

cineole

cineole

camphor

camphor

cineole

cineole

camphor

141

100
90
'Camphor' chemo-type

cumuative % frequency

80

n=208 from 6 trees GC confirmed


Globule Type 1. - many small

70
'Cineole' chemo-type

60

n=128 from 3 trees GC confirmed


Globule Type 2. - many large

50
40

Globule length at 70% cumulative frequency:

'Camphor' chemo-type

30

7.8 um for 'Camphor' chemo-type - Globule Type 1.


32.5um for 'Cineole' chemo-type - Globule Type 2.
19.8 um for 'Camphor' chemo-type - Globule Type 3.

n=527 from 10 trees GC confirmed


Globule Type 3 - Intermediate - small + large

20
10
0
1

13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85 89 93 97 101 105

oil globule length (m)

Figure 28. Cumulative percentage frequency for oil globule length in the leaves of
nineteen street trees of the two chemo-types of camphor laurel.

100

cumulative % frequency

90

Nimbin tree- -'Cineole' chemotype


n=53, 70% globules at 14.5 um

80

Globule Type 3.

70

Lismore tree -'Camphor' chemo-type


n=47, 70% globules at 4.8 um
Globule Type 1.

60
50

Uki tree - 'Cineole' chemo-type


n=84, 70% globules at 7.2 um

40

Comparison to GC of Street Trees, Figure 28:

Globule Type1.

Comparison to GC of Street Trees, Figure 4.2:


70% of the comparative globule lengths occur at:

30

7.8 um for 'Campho'r chemo-type - Globule Type 1.


32.5um for 'Cineole' chemo-type - Globule Type 2.
19.8 um for 'Camphor' chemo-type - Globule Type 3.

20
10
0

9 13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85 89 93 97 101 105 109 113

oil globule length (um)

Figure 29. Cumulative percentage frequency for oil globule length in the leaves of
three wild trees of the two chemo-types of camphor laurel.

142

3.4.3 Venation
Examination of the veins, veinlets and areolae of the leaf revealed no consistent visual
differences between the chemo-types.

3.4.4 Leaf Sclerophylly


The pilot sclerophylly test of the two chemo-types revealed a possible observable
relationship between leaf hardness and rigidity between the chemo-types. Ten of the
eleven camphor chemo-typed trees were found to have a sclerophyllous leaf form of
hard leathery leaves and thickened margins, while for the cineole chemo-type five of
the six trees had thin, pliable leaves with thin margins. Testing of leaf midrib
thickness using a micrometer revealed no significant difference in sclerophylly
between the chemo-types (F = 0.203; df = 1, 138; p = 0.653).

3.4.5 Olfactory Recognition


Following the pilot study using three olfactory trials it was determined that fine
cutting and placement of leaf material within a 12cm wide porcelain dish with a
volatile entrapping glass lid was the most efficient method for olfactory
differentiation. The use of plastic containers was found to taint the smell liberated
from the leaf, reducing successful olfactory identification. The resultant blind test of
the eighteen gas chromatographed trees using the porcelain container method (Table
15) identified an overall success rate of 90.6 % in the one hundred and eighty-one
blind tests examined. It was also demonstrated that olfactory recognition declined
from an average of 92.2 % during warm dry weather to 77% under cool wet
conditions. Age of the refrigerated leaf material was also found to affect the success
rate, which was on average 95% for one day old material, 91.3 % for two day old
material and 84.2 % for four day old material.

143

Table 15. The identification of the camphor laurel chemo-types using olfactory
recognition during blind testing of gas chromatographed leaves.

Test No.

Total

Age of
leaves
and
weather
conditions

1day old
warm
dry

2 day old
warm
dry

4 day old
warm
dry

1day old
warm
dry

2day old
warm
dry

1 day
old
warm
dry

2 day old
warm
dry

1 day old
warm
dry

1 day old
cool
wet

No. of
trees
sampled

18

16

19

18

21

21

19

23

26

181

No. of
Camphor
chemotypes

13

12

14

13

14

15

11

18

15

125

No. of
Cineole
chemotypes

11

56

No.
incorrect

18

Error
type

Cineole
as
Camphor

Cineole
as
Camphor

Cineole
as
Camphor

Cineole
as
Camphor

Cineole
as
Camphor

Nil

Cineole
as
Camphor

Cineole
as
Camphor.

Cineole
as
Camphor

Camphor
as
Cineole

%
Correct

94.4

93.8

84.2

94.4

90.5

100

89.5

91.3

77

90.6

3.5 Discussion
Bandulska (1928) described simple epidermal and internal morphology structures of
the camphor laurel leaf comparing it to three other Cinnamomum species including a
fossil Cinnamomum. In the study by Balasubramanian et al. (1993), petiole anatomy
was compared between four species of Cinnamomum including *C. camphora.
Although useful as a technique of differentiation between species this did not provide
information on possible variation below the species level, such as with chemo-type
studies.

144

Work by Santos (1930) on the genus Cinnamomum in the Philippines, provided


detailed descriptions and illustrations of the internal structure of the leaves and bark
which were compared between seven species, but not including *C. camphora.
However, the study was useful and informative as a guide in undertaking any
sectioning, staining and examination of *C. camphora sections. This is due to the
detailed nature of that morphological study, as well as there being much similarity in
cell structure within the Cinnamomum genus. This is apparent with the cellular
morphology of *C. zeylanicum demonstrated in the paper by Santos (1930) and *C.
camphora when compared to the sectioning work undertaken for this study.

At the taxonomic level of family, leaf and cuticle features of the Lauraceae have been
described in Australia by Christophel and Rowett (1996) for seven genera including
Cinnamomum, but not *C. camphora. This compared in detail the leaf features of
shape, venation, domatia, cuticle, stomata and trichomes between the genera of the
family, but not within the genera or species themselves.

The morphological studies of the camphor laurel leaf by Bandulska (1928), Santos
(1930), Balasubramanian et al. (1993) and Christophel and Rowett (1996) have not
been adequate for this current comparative study of the field differentiation of the
chemo-types. This is because they not only lack detailed morphological descriptions
of *C. camphora and, more importantly, they do not differentiate the chemo-types of
the tree. Overall the internal morphology and histology of *C. camphora has been
little studied, with the literature not able to provide any possible comparison of the
physical differences in the species below that level. In recognition, this phytotoxicity
research does warrant the further examination of morphological variation between the
two known chemo-types in New South Wales.

However, in this study of hundreds of leaf sections of both chemo-types using eleven
histological stains and repeated comparison of internal and external cellular
morphology and colour, the work revealed no consistent differences. A modified
parenchyma cell containing calcium oxalate, i.e. rhaphides as referred to by Bowes
(1996), was also seen to vary between the chemo-types in the initial work. However,
Santos (1930) described these as calcium oxalate crystals found in the genus

145

Cinnamomum, but staining with Pizzolato stain did not fully verify them to be
calcium oxalate as only a slight colour variation occurred. The most promising
approach involving the use of Sudan III stain appeared when the size and colour of the
oil globules of the secretory cells was initially assessed. In the final application of the
technique to the field population it was found to lack consistency between the chemotypes, identifying that oil globule size and colour is not dependant upon chemo-type
genetics.
Literature relating to solution or precipitate testing of *C. camphora is not known. In
the current study of chlorophyll clearing, plastid solution staining and solution
precipitation, no visual differences emerged. The reason for this may rest with a few
factors such as inadequate stains to target the volatile oils of the tree, age of the leaf
material influencing the volatile presence or composition, or the small amount of leaf
material used in the extraction.

Venation studies in Australia of the seven genera in the family Lauraceae including
the genus Cinnamomum by Christophel and Rowett (1996) describe differences in
veins, veinlets and areolae of the leaf, with *C. camphora not included in that study.
The pilot study revealed no visual differences in the venation features of the two
chemo-types. Therefore, further leaf clearing using nitric acid or other solvents was
considered not worthwhile.
References are not known relating to sclerophylly in camphor laurel. In consideration
of the broad geographical range of camphor laurel in its varying natural habitats in
Asia it would seem possible that sclerophyllous differences may well exist within the
species. This may be possible due the varying environmental conditions prevailing
across the natural range of these species, and the fact that discernable differences exist
in leaf morphology between the species of the genera as identified by Bandulska
(1928), Santos (1930), Balasubramanian et al. (1993) and Christophel and Rowett
(1996).

The initial chemo-typing of *C. camphora in Australia by Hirota and Hiroi (1967)
using olfactory recognition, identified two chemo-types which were later verified
through GC work by Stubbs and Brushett (2001) and Stubbs et al. (2004). The current

146

study revealed olfactory recognition to possess a high rate of success (average 95%
accuracy) under particular environmental and experimental conditions when gas
chromatographed leaves of the chemo-types were used as a reference aroma. The use
of plastics in the sealed plastic bag and the aspirator pilot tests tainted the volatile
aroma making differentiation difficult. This was overcome through the use of
porcelain and glass containers which not only entrap the volatiles from the finely cut
leaf material but do not taint the aroma. The age of the leaf in refrigerated storage was
also found to affect the success rate, presumably due to rapid volatile denaturing
within the leaf or volatile release soon after leaf harvesting. Success rates were
slightly diminished under cool, wet weather conditions due possibly to the lower
volatile release rates expected under such conditions.

During the gas chromatography undertaken for this study 18 additional terpenoids
were identified which have not be listed in the literature by Hirota and Hiroi (1967),
Chou et al. (1989), Stubbs and Brushett (2001) and Stubbs et al. (2004). Brushett
(pers. com. 2006) suggests that the differences shown between these studies are most
likely due to variation in oil synthesis of the tree as a result of seasonal fluctuations of
temperature, rainfall and the status of flowering and fruiting. This variability
demonstrates that terpenoids manufactured by the tree changes throughout the year
but with relatively little difference between trees within each chemo-type at any given
time. In addition, the synthesis of an extensive range of terpenoids indicates that a
complex allelopathy may occur apart from that originating from cineole and
camphor which are found at higher concentration.

3.5 Conclusion
Of the five tests applied to camphor laurel leaves to identify a consistent feature for
infield chemo-type differentiation, olfactory recognition was found to be the most
accurate, reproducible and inexpensive. An accuracy of between 91.3 and 100%
(average 95%) was achieved where leaf material was less than one day old, gathered
during warm dry weather, finely cut and placed in a 12cm wide porcelain bowl with
volatile entrapping glass lid, and compared against a GCd standard aroma. The trial
also demonstrated leaf material older than one day, as well as wet cool weather, does

147

influence the efficiency of the technique. This reduces accuracy to between 77 and
93.8% (average 87%).

Overall the average success rate irrespective of time or

weather conditions equates to an accuracy of 90.6%.

The other techniques trialed such as sectioning and staining, leaf solution clearing and
precipitate comparison, venation and sclerophylly were found to possess no consistent
visual or measured differences between the chemo-types.

This developed olfactory technique can now be applied to a large number of camphor
laurel trees rapidly, inexpensively and reliably which enables the study of seedling
growth below each chemo-type and other vegetation types to identify if phytotoxic
effect is active in the field situation.

148

-Chapter 4:Field Assessment of Camphor Laurel Allelopathy


in Vascular Plant Seedlings.
4.1 Introduction
The allelopathy glasshouse trial (Chapter 2), examined camphor laurel extract
influences on germination and growth of 52 species of vascular plant seedlings and 27
species of soil algae present in the vegetation assemblages of north-eastern New
South Wales. For the assessment of allelopathy under field conditions both these plant
groups present some difficulty for the measurement of plant growth.

Soil algae in particular would be less conducive for field measurement due to their
microscopic size and growth variability through time and space in the soil profile.
This results from fluctuations in temperature, moisture and light throughout the year
and its rapid influence on the growth of these organisms. Therefore, possible
identification of camphor laurel allelopathy under field conditions required a group of
plants which possessed easily measurable growth characteristics and greater resilience
to the micro-climate fluctuations experienced in the soil profile than do the algae.

Vascular plant seedlings in this case were a more appropriate plant group to use due to
their robust size and greater resilience to changes in microclimate.

They were

important to assess as seedling recruitment and growth in vascular plant communities


are intrinsic to successional change over time. The use of the same vascular species
tested in the glasshouse trial for the field comparisons would also provide a level of
allelopathic comparison. Additionally, the use of species with a highly significant
response to the camphor laurel leachates in the glasshouse trial would be more likely
to demonstrate allelopathic effect in the field. This provided a selected targeted
species group for the field comparison of effect of both the cineole and camphor
chemo-types.

149

The vascular plant glasshouse trial measured germination over time and growth as
shoot length, root length, leaf number and leaf area. Of these, the growth of shoot
length, leaf number and leaf area were identified as easily measurable in the field.
Root length however, would be more difficult to measure due to problems of soil root
removal without breakage, particularly from highly root compacted soils. This
enabled a robust comparison of growth between the glasshouse trial and field
measurements for shoot length, leaf area and leaf number.

Importantly, a number of biotic and abiotic factors are responsible for inhibitory or
stimulatory growth responses in vascular plant seedlings due to factors other than
allelopathy. These may include variation in light, soil moisture, nutrients and pH;
plant competition above and below the ground-surface, seedling age, animal browsing
and genetic variation inherent within seeds (Whittaker 1970, Rice 1984). Whittaker
(1970) identifies that some of these factors, particularly high shade and root
competition for nutrients and water has the effect of increasing the influence of
allelopathy. Therefore, the design of the field sampling aimed to accommodate as
many of these variables as was practical to minimize growth influences resulting from
non-allelopathic factors. In other words, the comparison of plant growth should be
from sites that demonstrate relative environmental consistency so that any possible
allelopathic growth influences are more likely to be shown without the complexity
resulting from environmental variability in the landscape.

4.2 Aim and Hypothesis


The aim of the field assessment was to establish if the allelopathic responses
identified in the growth of the most responsive vascular plant seedlings from the
glasshouse trial:

demonstrate allelopathic effect below the camphor laurel canopy in


north-eastern New South Wales, and

whether significant differences exist in allelopathic effect between the


cineole and camphor chemo-types of camphor laurel on shoot
length, leaf area and leaf number in the field.

150

The null hypothesis H0 being tested was:

there is no significant allelopathic effect below the camphor laurel


canopy on vascular seedling shoot length, leaf number and leaf area of
species identified in the glasshouse trial which displayed the most
significant responses to camphor laurel leachates, and

there is no significant difference in allelopathic effect on seedling


growth of these species for shoot length, leaf number and leaf area
between the cineole and camphor chemo-types of camphor laurel
under field conditions.

4.3 Methods
For the purposes of this study a seedling was defined as a vascular plant 15cm or less
in height. The target vascular plant seedlings for the field sampling identified from
the allelopathy glasshouse trial which displayed a significant influence of the camphor
laurel leachates on three or four of the growth parameters of shoot length, root length,
leaf number and leaf area of seedlings included for the cineole chemo-type: A.
caerulea, C. bartramia, *L. lucidum, L. confertus, O. aemulus, C. hypoglauca, E.
intermedia, E. saligna, G. semiglauca, H. flavum, L. longifolia, M. tanarius, *P.
clandestinum, S. capsicoides, S. luehmannii, S. paniculatum, V. hederacea, and for the
camphor chemo-type: C. hypoglauca, C. viminalis, C. enervis, F. coronata, L.
longifolia, L. confertus, S. paniculatum, V. hederacea, *C. grandiflorum, C.
bartramia, E. saligna, *L. lucidum, O. aemulus, *P. clandestinum, S. capsicoides and
T. australis. Apart from these species it was also considered important to measure the
seedlings of *C. camphora even though the glasshouse trial identified that an error
had occurred in the control of the fresh seed, as it would provide an understanding
of how camphor laurel seedlings respond to its own allelochemicals in the field.
Seedling identification used pressed specimens from the glasshouse trial for positive
verification.

Neilan (2004) found that seedling diversity under camphor laurel increased the closer
to larger rainforest areas of the Nightcap range. For this reason, field locations were
151

identified close to these rainforest seed source areas to provide adequate seedling
numbers, species and vegetation variability for comparison. The study sites were
located on ridge-tops at Rosebank (S28.69310E153.39404) and Lismore
(S28.78450E153.24353), and riparian areas at Nimbin (S28.58624E153.21686)
and Uki (S28.40752E153.33607).
Sites for the 1m2 quadrat sampling were selected below camphor laurel trees of both
chemo-types determined by olfactory technique developed in Chapter 3 where
present, and other vegetation as the control using stratified random sampling where
target seedlings were identified. Where possible a minimum of 3 sites and 3 quadrat
replications per site was adopted for each target species under each of 3 canopy
types i.e. the cineole and camphor chemo-types of camphor laurel and control
vegetation other than camphor laurel. Sampling was conducted for a period of 12
months during both winter and summer across a variety of sites of varying soil type
and slope. The technique recorded the presence of the target species in each quadrat.
Biotic and abiotic variables were kept as homogenous as possible between the sites
and quadrats. The criteria used to establish sampling consistency at site included
foliage projective cover of 60%, similar slope, aspect, elevation, soil type and pH,
litter depth and spread, root competition and disturbance. Foliage projective cover
(FPC) was calculated as a percentage taken from 50 points per site measured with a
foliage projective tube; slope as a percentage measured by clinometer; aspect by
compass; elevation and site location by GPS; soil order as described by McDonald et
al. (1990), colour by Munsell (2000) and pH using barium sulphate and indicator dye;
litter depth (mm); litter spread as a percentage coverage of quadrat i.e. 100-60%, 6040%, < 40% and 0%; root competition as feeder roots absent, feeder roots highly
compacted or feeder roots not highly compacted. All target seedlings below 15cm in
height in the quadrats were labeled, pressed and mounted. Growth measurements were
later recorded for each seedling which included shoot length (mm), leaf number, and
leaf area as length and width (mm) of the largest leaflet.

With *C. camphora seedlings 3 sites at Rosebank, Nimbin and Uki were sampled
using 12 quadrats for the control with 356 seedlings, 14 quadrats for the camphor
with 1004 seedlings and 6 quadrats for the cineole with 360 seedlings. In the case of
*L. lucidum, 2 sites located at Rosebank and Lismore resulted in 13 quadrats for the
152

control with 895 seedlings and 9 quadrats for the camphor canopy with 677
seedlings. G. semiglauca seedlings were measured at 2 sites located at Rosebank and
Uki which produced 12 quadrats for the control with 128 seedlings and 9 quadrats
for the camphor with 199 seedlings. The forest grass O. aemulus was sampled as a
mature plant with the largest inter-node length per 15cm of stolon measured as a
substitute for shoot length. The number of leaves was also counted and the largest leaf
area measured for the 15cm stolon length. The grass was only sampled in a single site
at Uki producing 3 quadrats for the control with 36 stolons measured and 3 for the
camphor with 45 stolons.

The camphor laurel chemo-types were assessed using olfactory recognition


developed in Chapter 4, enabling a large number of trees to be assessed rapidly,
reliably and at little expense. This involved daily leaf gathering from known gaschromatographed camphor laurel trees of the cineole and camphor chemo-types
located at Uki, Nimbin and Lismore. Cutting and placement of leaf material of the
GCd tree and unknown tree in separate ceramic bowls with enclosing glass lids
enabled concentration of the volatiles without contamination, and comparison of the
distinctive odor to the GCd material.

Statistical analysis of shoot length, leaf number and leaf area used a two-sample t-test
assuming equal variance between the control and the cineole treatment, the
control and the camphor treatment and the cineole and the camphor treatment
where these chemo-types were present. Comparisons were then made between the
effect of the control and the two treatments and the difference between the
treatments.

4.4 Results
Seedlings of *C. camphora, *L. lucidum, G. semiglauca and O. aemulus were found
in large enough numbers across a variety of sites to enable quadrating. Difficulty was
experienced in finding adequate numbers of camphor laurel trees of the cineole
chemo-type in bush-land which possessed target seedlings below their canopy. It
was found that the camphor chemo-type occupies both dry ridge-tops as well as
moist riparian locations while the cineole chemo-type was found exclusively in
153

moist riparian habitat and was relatively uncommon with an approximate ratio mix
with the camphor chemo-type of 1:10.

4.4.1 Sites, Quadrats and Sample Size


The environmental attributes recorded for each site and quadrat for the study appears
in Appendix 14; Table A14.1 to A14.8. Comparisons of quadrat variables at each site
and between sites identified relative environmental consistency at site as well site
variability between the sites. The variability in environmental attributes recorded
across all sites were FPC: 60-90%, Slope: 0-12, Aspect: north, south, east, north-east
and flat, Elevation: 4.3-60m, Soil Orders: Ferrasol and Podosol, Soil Colour: 2.5 YR
2.5/(1, 3, 4), 10YR 2/(1, 2) and 7.5YR 3/2, Soil pH: 5.5-7.2, Litter Depth: 20-60mm,
Litter Spread: 40-100%, Root Zone: highly compacted with a few not compacted, and
Disturbance: animal browsing absent and roadside mowing, river wash and no
disturbance occurring at some sites.

4.4.2 Seedling Measurements


Shoot lengths, leaf numbers and leaf areas were grouped together across the sites for
analysis and comparison of effect so as to reduce variation due to any distinct
environmental site attributes.

Sorted seedling shoot lengths, leaf numbers and leaf areas of *C. camphora across the
sites at Rosebank, Nimbin and Uki under the camphor and cineole chemo-types
and other vegetation as the control are shown in Figure 30a to c.

For the seedling shoot lengths of *C. camphora (Figure 30a), minimal measurements
started at 43mm under the control, 25mm for the camphor and 40 mm for the
cineole canopy with 150mm recorded as the maximal length attained for each of the
treatments. However, a reduction was evident in seedling lengths under the camphor
canopy with these being below the control when seedlings reached a height of
50mm while the cineole canopy rose above the control commencing at 60mm. In
the case of leaf numbers for the species at this site (Figure 30b), these began at 1 for
the control, camphor and cineole canopy. The maximal leaf numbers attained in
the control were 26, the camphor 12 and cineole 11. A reduction was evident in
leaf numbers under the camphor and cineole canopy compared with the control,
154

when leaf number exceeded 2 for camphor and 5 for cineole. For leaf areas (Figure
30c), the control had minima of 6mm2, camphor of 21mm2 and cineole of
50mm2. The maximal leaf areas obtained for the control were 4400mm2, camphor
3080mm2 and cineole 1988mm2. Comparisons between the leaf areas under both the
cineole and camphor canopies and the control demonstrated that leaf area was
reduced for seedlings under these canopies. This reduction in leaf area commenced at
600mm2 for the cineole and 6 mm2 for camphor.

(a).
160

n = 356

n = 1004

140

n = 360
Sorted shoot length sizes
(mm)

120
100
80
Control
60

Camphor
Cineole

40
20
0
0

200

400

600
800
Number of seedlings measured

1000

1200

(b).
60

Sorted leaf numbers per seedling

50

40

30

n = 1004

n = 356

Control
Camphor

20

Cineole

n = 360

10

0
0

200

400

600
800
Number of seedlings measured

155

1000

1200

(c).
5000

n = 356

Sorted leaf area of largest leaf (mm)

4500
4000
3500

n = 1004

3000
2500

n = 360
2000
1500

Control
Camphor

1000

Cineole

500
0
0

200

400

600
800
Number of seedlings measured

1000

1200

Figure 30. Sorted measurements of *C. camphora seedlings below the


chemo-types of *C. camphora and 'control' vegetation across all sites
sampled for: (a) shoot lengths; (b) leaf numbers; (c) leaf areas.

For *L. lucidum the sorted shoot lengths, leaf numbers and leaf areas of seedlings
under the camphor chemo-type and the control at Rosebank and Lismore are
shown in Figure 31a to c.

With shoot lengths of *L. lucidum recorded across the sites (Figure 31a), the control
commenced with a length of 20mm and the camphor with 18mm. The longest length
recorded for both the control and camphor was 150mm. An increase in shoot
length above the control for seedlings under the camphor canopy was recorded,
commencing when seedlings were 30mm high. For the leaf numbers of the species
(Figure 31b), these were minimal at 1 under the control and camphor canopy and
completing with 54 for the control and 31 for camphor. Leaf numbers became
greater than the control for the seedlings under the camphor chemo-type compared
with the control when leaf numbers exceeded 7. In Figure 31c leaf areas of the
largest leaf had a minimal of 11mm2 for the control and 15mm2 under the camphor
chemo-type. The maximal leaf areas recorded was 2860mm2 for the control and
1690mm2 for camphor. An increase above the control occurred for the camphor
canopy at 250mm2.

156

(a).
160

n = 677

n = 895

140

Sorted shoot lengths


(mm)

120
100
80
60

Control
Camphor

40
20
0
0

200

400

600
800
Number of seedlings measured

1000

1200

(b).
60

n = 895

Sorted leaf numbers per seedling

50

40

n = 677
30

Control

20

Camphor
10

0
0

200

400
600
Number of seedlings measured

800

1000

1200

(c).
5000

Sorted leaf area of largest leaf


(mm)

4500
4000
3500

n = 895

3000
2500

n = 677

2000
1500
1000
Control

500

Camphor

0
0

200

400

600

800

1000

Number of seedlings measured

Figure 31. Sorted measurements of *L. lucidum seedlings below


the 'camphor' chemo-type of *C. camphora and 'control'
vegetation for all sites sampled for: (a) shoot lengths; (b) leaf
numbers ; (c) leaf areas.

157

1200

The sorted shoot lengths, leaf numbers and leaf areas of G. semiglauca seedlings
under the camphor chemo-type and the control at Rosebank and Uki appear in
Figure 32a to c.

With sorted shoot lengths of G. semiglauca plotted across the sites (Figure 32a), the
control had a minimal length of 20mm and the camphor with 40mm. Both the
control and camphor attained a maximal length of 150mm. Shoot length was less
under the camphor compared to the control canopy, commencing when seedlings
were 30mm in height. With leaf numbers for G. semiglauca (Figure 32b), minimal
measurements started at 3 under the control and 2 under the camphor canopy with
12 being the maximal recorded for the control and 9 for camphor. Leaf numbers
were fewer below the control for camphor when leaf numbers exceeded 6. Leaf
areas for this species (Figure 32c) demonstrated that the largest leaf area commenced
at 24mm2 for the control and 2mm2 under the camphor canopy. Maximal areas of
900mm2 for the control and 600mm2 for camphor were found. Leaf area was less
below the control than the camphor canopy being apparent from the
commencement of measurements.

(a).
160

n = 128
n = 199

Sorted shoot lengths (mm)

140
120
100
80
60

Control
Camphor

40
20
0
0

200

400
600
800
Number of seedlings measured

158

1000

1200

(b).

Sorted leaf numbers per seedling

60

50

40

30

20

Control

n = 128
n = 199

Camphor

10

0
0

200

400

600

800

1000

1200

Number of seedlings measured

(c).
5000

Sorted leaf area of largest leaf


(mm)

4500
4000
3500
3000
2500
2000
1500

n = 128
Control

n = 199

1000

Camphor

500
0
0

200

400

600
800
Number of seedlings measured

1000

1200

Figure 32. Sorted measurements of G. semiglauca seedlings below the


'camphor' chemo-type of *C. camphora and 'control' vegetation across all
sites sampled for: (a) shoot lengths; (b) leaf numbers; (c) leaf areas.

Figure 33a to c, displays the sorted inter-node lengths, leaf numbers and leaf areas of
O. aemulus seedlings under the camphor chemo-type and the control at Uki.

With the inter-node lengths of O. aemulus that were recorded across the sites (Figure
33a), the control had a minimal length of 10mm and the camphor with 14mm. The
maximal was 30mm for the control and 40mm for camphor. An increase in the
inter-node length of camphor above the control was noted being apparent from the
159

commencement of measurements. With leaf numbers (Figure 33b), the minimal was 6
for the control and 5 for the camphor canopy. The maximal leaf numbers were 13
for the control and 12 for camphor. Leaf numbers appeared slightly depressed
under the camphor canopy when compared with the control. This commenced
from the minimal measurements until 10mm was reached when it became similar to
the control. Mimimal leaf areas of the largest leaf (Figure 33c), was 48mm2 for the
control and 70mm2 under the camphor canopy. The maximal leaf areas recorded
were 250mm2 for the control and 280mm2 for the camphor canopy. Leaf area rose
slightly above the control for the camphor canopy and was apparent from the
beginning of measurements.

(a).
160

Sorted inter-node lengths


(mm)

140
120
100

Control

80

Camphor

n = 36

60

n = 45

40
20
0
0

10

15
20
25
30
Number of stolons measured

35

40

45

50

(b).

Sorted leaf numbers per stolon

60
50
40
Control

30

Camphor

20

n = 45

n = 36

10
0
0

10

15

20

25

30

Number of stolons measured

160

35

40

45

50

(c).
5000

Sorted leaf area of largest leaf


(mm)

4500
4000
3500
3000
2500
Control

2000

Camphor

1500
1000

n = 45

n = 36

500
0
0

10

15

20

25

30

35

40

45

50

Number of stolons measured

Figure 33. Sorted measurements per 150mm of stolon for O. aemulus below the
'camphor' chemo-type of *C. camphora and 'control' vegetation for: (a) internode lengths; (b) leaf numbers; (c) leaf areas.

The sorted shoot and inter-node lengths, leaf numbers and leaf areas of all seedlings
other than *C. camphora i.e. *L. lucidum. G. semiglauca and O. aemulus are shown
collectively in Figure 34a to c. These were measured under the camphor chemo-type
and the control across all sites sampled at Rosebank, Lismore, Nimbin and Uki.

With the sorted shoot and inter-node lengths of the combined species recorded across
the sites (Figure 34a), the control measurements had a minimal length of 10mm and
the camphor with 14mm. The maximal length attained was 150mm for both the
control and camphor. The camphor was greater than the control between 1475mm, less at 75-100mm and greater at 100-150mm. This resulted in a fluctuation
which was visually difficult to determine direction of effect for these combined
species until the largest sizes were reached at 100mm where the camphor canopy
demonstrated reduced lengths. In the case of leaf numbers in Figure 34b, the minimal
was 1 for both the control and camphor reaching a maximal of 54 for the control
and 31 for the camphor canopy. The leaf numbers of both the control and
camphor fluctuated regularly across each other on the graph making visual
observation of effect difficult to establish. However, the maximal leaf number reached
was markedly lower under the camphor canopy. For the areas of the largest leaf
shown in Figure 34c, measurements began at 11mm2 for the control and 2mm2
161

under the camphor canopy resulting in a final area of 2860mm2 for the control and
1690mm2 for the camphor canopy. Similar to leaf number, leaf areas of both the
control and camphor regularly fluctuated across each other but with the largest leaf
area obtained markedly reduced under the camphor canopy.

(a).
160
n = 912

Sorted shoot & internode lengths


(mm)

140
n = 1068
120
100
80
60
40
Control

20

Camphor

0
0

200

400
600
800
Number of seedlings measured

1000

1200

(b).

60

Sorted leaf numbers per seedling

n = 1068
50
40
n = 912
30
20

Control
Camphor

10
0
0

200

400
600
800
Number of seedlings measured

162

1000

1200

(c).

5000

Sorted leaf area of largest leaf


(mm)

4500

n = 1068

4000
3500
3000

n = 912

2500
2000
1500
1000
Control

500

Camphor

0
0

200

400
600
800
Number of seedlings sampled

1000

1200

Figure 34. Sorted measurements of all sampled species other than *C.
camphora found below the 'camphor' chemo-type of *C. camphora and
'control' vegetation across all sites for: (a) shoot lengths; (b) leaf numbers;
(c) leaf areas.

4.4.3 Significant Effects below the Canopy


The t-tests for the field effects of the camphor laurel chemo-types on shoot length,
leaf number and leaf areas for each species across the sites are shown in Appendix 15.

With *C. camphora seedlings found below the camphor chemo-type the t-tests
demonstrated that no significant difference (p = 0.148) occurred for mean shoot length
with 107mm for the control and 105mm for camphor, while under the cineole
chemo-type the mean length significantly increased (p = 4.801-19) from 107mm in the
control to 122mm under the cineole. A significant difference (p = 2.239-36) was
also apparent between camphor with a mean length of 105mm and cineole with
122mm. For leaf numbers the t-tests of *C. camphora seedlings identified a
significant difference (p = 1.783-81) between the control and camphor with the
mean significantly reduced from 7 to 4 leaves respectively. In the case of the control
and cineole the mean leaf number was also significantly reduced (p = 2.36-14) from 7
to 5 respectively. A significant difference (p = 1.039-35) was also noted in the
comparison of cineole with a mean of 4 and camphor with a mean of 5 leaves. The
t-tests for the leaf areas of *C. camphora seedlings revealed that a significant
difference (p = 2.065-8) existed between the control with a mean of 824mm and
163

camphor with 676mm, identifying a significant reduction under the camphor


canopy. A significant difference (p = 0.005) was also noted for the cineole canopy
where the mean leaf area of 722mm was less than the control which was 824mm.
With the comparison of the cineole and camphor leaf areas a significant difference
(p = 0.031)also emerged where the camphor mean of 676mm was below the
cineole at 722mm, identifying that the cineole chemo-type results in a higher leaf
area.

For the species *L. lucidum the t-tests demonstrated a significant difference in shoot
length (p = 0.001), leaf number (p = 1.026-11) and leaf area (p = 6.593-5) under the
camphor canopy compared to the control. With shoot length the mean was
significantly reduced from 70mm in the control to 66mm under the camphor
canopy. In the case of mean leaf number this was significantly reduced from 10 for
the control to 8 leaves for the camphor canopy. Similarly, mean leaf area was
significantly reduced from 284mm for the control to 235mm under the camphor
canopy.

In the species G. semiglauca, t-test comparisons identified a significant reduction in


shoot length (p = 5.92-24), leaf number (p = 1.704-9) and leaf area (p = 2.434-16) when
the control and camphor canopy were compared. For shoot length this was
significantly reduced from a mean of 95mm in the control to 71mm for the
camphor canopy. Leaf numbers in the control of 6 was significantly reduced to 5
for the camphor, and with leaf area the significant reduction was from 257mm for
the control to 123mm under the camphor canopy.

With O. aemulus sampled at only 1 site the t-tests revealed that the mean inter-node
length significantly increased (p = 1.364-9) below the camphor canopy from 19mm
for the control to 28mm. For leaf number, this significantly decreased (p = 1.313-7)
from a mean of 9 in the control to 7 leaves for the camphor canopy. However, in
the case of leaf area the mean significantly increased (p = 1.8-5) from 104mm in the
control to 154mm for the camphor canopy.

164

With the combined species over all sites and excluding *C. camphora, a significant
reduction in shoot and inter-node length (p = 8.14-6), leaf number (p = 5.836-19) and
leaf area (p = 1.44-10) was identified by the t-tests. This resulted in a significant
reduction in combined mean shoot and inter-node length from 72mm for the control
to 66mm for the camphor, a significant reduction in mean leaf number of 9 in the
control to 7 under camphor, and a significant reduction in mean leaf area of
273mm in the control to 207mm under the camphor canopy.

4.5 Discussion
In Taiwan, the research by Chou et al. (1989) into the effects of camphor laurel
allelochemicals on the growth of vascular plant species in a glasshouse trial
demonstrated reductions in many growth parameters. These included seedling length
and fresh weight of the grass *Miscanthus floridulus; root initiation of Kikuyu
(*Pennisetum clandestinum) and radicle growth of perennial ryegrass (*Lolium
perenne), annual ryegrass (*Lolium multiflorum) and Chinese cabbage (*Brassica
oleracea). Camphor allelopathy was attributed to have increasing effect in reducing
growth as extract and leachate percentages increased in the treatments. No studies are
known where camphor laurel allelopathy was directly measured in the field. However,
Paul et al. (2010) found in a glasshouse study that soil from below camphor laurel
reduced the seedling growth rates by 25% for Alphitonia excelsa, Guioa semiglauca
and Omalanthus nutans being common rainforest regeneration species in northeastern NSW.

With the allelopathy glasshouse trial for this research (Chapter 2), 46 species of
vascular plants from north-eastern NSW were found to possess significant growth
reductions due to the effect of the leachates from the cineole and camphor chemotypes in 86% of species for shoot length, 64% for radicle length, 54% for leaf number
and 71% for leaf area with little difference between the effects of the two chemotypes. Some species did display growth increases or no significant effect further
identifying that the allelopathic influence can vary as a stimulation, a reduction or as
no significant influence.

165

However, before glasshouse data can be used to identify field effect, seedlings in the
natural environment needed to be assessed for verification of allelopathic influence as
the complex interactive variables associated with the natural environment do
influence plant growth and must be considered.

With the 4 species i.e. *C. camphora, *L. lucidum, G. semiglauca and O. aemulus
assessed for this field study of seedling growth under camphor laurel and other
canopy types, significant growth reductions mostly occurred as well as some
significant growth increases across the parameters measured.

In *C. camphora the canopy of the camphor chemo-type had no effect on mean
shoot length of its own seedlings with only a 2% reduction, while under the cineole
canopy a significant stimulation increased mean shoot length by 14.4% compared to
the control. There was also a significant difference between the mean shoot length
under the cineole compared to the camphor canopy with a 16.7% increase for
cineole. This indicates that under field conditions this chemo-type has greater
influence on shoot growth of its own species. For mean leaf number of the camphor
laurel seedlings a significant reduction occurred under the camphor canopy of
39.3% and the cineole canopy of 22.9% compared to the control. Statistical
comparisons between the cineole and camphor identified that the seedlings under
the cineole canopy had mean leaf numbers significantly greater than the camphor
canopy by 27%. This indicated that the cineole chemo-type has greater effect than
the camphor chemo-type on influencing leaf numbers of its own species. For mean
leaf area of both the camphor and cineole canopies these were significantly
reduced for the camphor by 18% and the cineole by 12.3%. Statistical comparisons
between the mean leaf areas under cineole and camphor showed that a significant
difference existed between these two chemo-types with a 6.9% increase of the
cineole above the camphor indicated that the cineole has greater effect at
influencing leaf area than the camphor canopy for its own species. Even though no
effect on mean shoot length occurred below the camphor chemo-type and a
stimulation was found under the cineole, the trend towards reduction in leaf numbers
and leaf areas indicates that both camphor laurel chemo-types reduce the growth of its
own seedlings. Comparisons of these field data for camphor laurel to the glasshouse
allelopathy trial of this species for germination (Appendix 1, Figure A1.17), the
166

deviance test for germination (Appendix 9) and the t-tests of growth (Appendix 11),
did not verify the direction of these growth reductions even though germination for
both chemo-types was below the control in the glasshouse. The reason for this could
lie with the fresh seed trialed for camphor laurel which failed due to a desiccation
event in the control and seed had to be used which originated from the soil profile.
Furthermore, the field environment is also substantially more complex than a
glasshouse and other variables such as root and canopy competition were absent in the
glasshouse.

In the case of *L. lucidum, the statistics revealed that mean shoot length, leaf number
and leaf area were significantly reduced below the control for the camphor canopy.
With mean shoot length the significant reduction was 6.1%, with mean leaf number it
was 19.4% and with mean leaf area 17.4% when compared to the control. With the
comparison of this data to the glasshouse allelopathy trial for germination (Appendix
1, Figure A1.13) the deviance test for germination (Appendix 9) and the t-tests of
growth (Appendix 11) verification of allelopathic effect as a growth reduction was
identified for shoot length, leaf number and leaf area for the camphor chemo-type.

For G. semiglauca statistical comparisons identified a similar trend to *L. lucidum but
more acute with mean shoot length, leaf number and leaf area significantly reduced
for the camphor below the control canopy. This was found to be 25.2% for mean
shoot length, 15.4% for mean leaf number and 51.6% for mean leaf area when
compared with the control. In the comparison of this with the glasshouse data
(Appendix 1, Figure A1.6), the deviance test for germination (Appendix 9) and the ttests of growth (Appendix 11) a reduction for leaf area only was verified in the
camphor chemo-type.
Only 1 site was measured for O. aemulus with mean inter-node length significantly
increased above the control by 49.7%, significantly reduced by 25.8% for mean leaf
number and significantly increased by 48% for mean leaf area. This variation
indicates that although lengthening of the inter-nodes occurs with lower leaf numbers
as a result, the leaf area actually increased above the control. When this species was
compared with the germination and growth analysis of the glasshouse data (Appendix
1, Figure A3.1) the deviance test for germination (Appendix 9) and the t-tests of
167

growth (Appendix 11) it revealed an opposite influence on shoot growth, leaf number
and leaf area. However this should be viewed with caution due to low sample size
with no site replication.

When the species are viewed in combination but with the exclusion of *C. camphora
i.e. *L. lucidum, G. semiglauca and O. aemulus, the general allelopathic trend was
identified for the sampled assemblage. In the statistical comparisons the collective
mean shoot length was significantly reduced by 7.9%, leaf number by 22.3% and leaf
area by 24.1%, which clearly establishes that allelopathy is active in the sampled
assemblage, resulting in a general trend towards growth reduction. These reductions
in growth are similar to those found by Paul et al. (2010) during a glasshouse trial
using soil from below camphor laurel. That study attributed the growth reductions to
physical and biochemical changes in the soil below camphor laurel.

The comparisons of mean shoot length and leaf area influenced by the camphor
canopy across the species tested (Appendix 15), also identifies that camphor laurel has
a longer mean shoot length and larger mean leaf area than other seedlings measured.
Even though growth reduction was indicated to be active for shoot length, leaf
number and leaf area across the seedling species measured in the field, including *C.
camphora, this species height and leaf area dominance would provide for substantial
growth above many other species. What this suggests is that allelopathy in
combination with the seedlings high growth attributes under such conditions provides
a competitive advantage for camphor laurel.

When the significant growth effect obtained from this field trial for each species is
compared with the glasshouse growth data in Chapter 2 it identifies that effect is
greater in the field than in the glasshouse. What this suggests is that high root and
light competition imposed by the camphor laurel tree and possibly interacting
environmental factors appear to exacerbate the allelopathic effect, as identified by
Whittaker (1970) to generally occur with allelopathy.

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4.6 Conclusion
Olfactory recognition of the camphor laurel chemo-types in the field trial which were
developed in Chapter 3 revealed that the camphor chemo-type occurs on ridge-tops
and riparian areas while the cineole chemo-type was found only in riparian areas in
an approximate ratio of 1 cineole to every 10 camphor trees. This resulted in a
paucity of cineole trees for seedling measurement under that chemo-type with
camphor laurel seedlings only being compared. What this also suggests is that the two
chemo-types may be distinct in their ability to withstand soil dryness with the
camphor chemo-type being more resilient to the drier ridgeline conditions and could
possibly explain the paucity of the cineole trees and possible differences in effect
between the chemo-types other than allelopathy. This may further indicate that the
camphor chemo-type may possess dispersal, recruitment or competitive attributes
which may make it more competitive.

The field effects of camphor laurel allelopathy due to the camphor canopy were
identified as a significant reduction in mean shoot length for *L. lucidum, G.
semiglauca and the sampled assemblage, in mean leaf number for *C. camphora, *L.
lucidum, G. semiglauca, O. aemulus and the sampled assemblage, and mean leaf area
for *L. lucidum, G. semiglauca and the sampled assemblage. With the camphor
chemo-type a significant growth increase was recorded for mean inter-node length
and leaf area in O. aemulus and no effect on shoot length for *C. camphora was
found. With the cineole canopy of camphor laurel it was found that the mean shoot
length of *C. camphora seedlings was significantly increased while mean leaf number
and leaf area were significantly reduced.

Statistical comparisons of seedling shoot length, leaf number and leaf area between
the cineole and camphor canopies also revealed that camphor canopy has a
greater influence than the cineole at reducing growth across the measured
parameters for its own seed.

The field study comparison to the glasshouse trial further identified that even though
camphor laurel allelopathy is active on its own seedlings these maintain competitive
advantage through possessing higher shoot length and leaf area than the other species
169

assessed suggesting that allelopathy in combination with high competitive growth of


plant is active. The comparison between the glasshouse trial and field assessment of
allelopathic effect may also suggest that in the field situation it may be substantially
higher. Factors such as root and canopy competition and possibly environmental
variables as indicated for some allelopathic situations by Whittaker (1970) may be
responsible for the observed exacerbation although this is speculative.

The environmental variables recorded for each site and quadrat for the study does
identify relative site consistency being applicable for statistical comparison.
Inconsistencies found between the field analysis and the glasshouse analysis are
possibly attributed to lack of fresh seed for *C. camphora and low sample size and
site replication for O. aemulus. However for *L. lucidum significant reductions were
recorded for field shoot length, leaf number and leaf area which were similar with the
glasshouse growth trial. Similarly, G. semiglauca had reduced mean leaf area in the
field situation which was also similar with the glasshouse findings.

The summary of the findings which address the hypothesis are:

there is a significant allelopathic effect below the camphor laurel


canopy on vascular seedling shoot length, leaf number and leaf area of
species identified in the glasshouse trial which displayed the most
significant responses to camphor laurel leachates, and

significant difference in allelopathic effect on seedling growth of these


species for shoot length, leaf number and leaf area between the
cineole and camphor chemo-types of camphor laurel under field
conditions was not possible to assess for species other than camphor
laurel. This was due to the rarity of the cineole chemo-type.
However, seedling shoot length, leaf number and leaf area of camphor
laurel does significantly vary between the cineole and camphor
canopies. A greater influence on these parameters below the camphor
canopy was apparent.

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- Chapter 5:Camphor Laurel: Potential and Possibilities.

5.1 Outcomes of the Phytochemistry and Allelopathy Research


The hypothesis tested for the glasshouse trial identified that a high number of common
native plants, woody weeds and soil algae were significantly influenced by the extracts of
both the cineole and camphor chemo-types of camphor laurel. With vascular plants,
phytotoxicity resulted in a significant influence in the time taken to germinate with the
occurrence of both stimulation and inhibition depending on the species affected. No
significant difference between the effects of the chemo-types was found for the time
taken to germinate for whole assemblage tested.

With the growth of seedlings,

phytotoxicity significantly inhibited radicle length, shoot length, leaf number and leaf
area in many species with some variation depending on chemo-type. For soil algae, the
phytotoxicity resulted in a significant delay taken in the time required to cover the soil
and a significant reduction in diversity of species with no significant difference between
the chemo-types. These glasshouse findings indicate that phytotoxicity resulting from
camphor laurel extracts has potential to be active in the field situation.
For the hypothesis tested in the field trial it was found that the growth parameters of shoot
length, leaf area and leaf number were significantly reduced below the camphor chemotype of camphor laurel. No assessment of the cineole chemo-type was possible except
with camphor laurel seedlings due to the rarity of that chemo-type in the wild. However,
with camphor laurel seedlings the camphor canopy was identified to have a significantly
greater influence than the cineole chemo-type at reducing growth of shoot length, leaf
area and leaf number.
The key outcomes from the research that might be useful in management are that
leachates originating from camphor laurel do influence vascular plant succession and
hence the long-term composition of regenerating plant assemblages which may place
stress upon less robust species and result in a less diverse assemblage. Soil algae are also
dramatically influenced with slower growth rates and a dramatic loss of diversity which
suggests that soil processes may also be affected such as soil structure, wetability, food
171

web relationships and germination ability of seeds. This identifies that both a direct
allelopathy is resulting from chemical release into the upper soil horizon is occurring and
that an indirect allelopathy may also result due to loss of soil algal benefits. This
phytotoxic influence alone now identifies camphor laurel to be of greater threat to the
long term maintenance of plant biodiversity in the Big Scrub region than previously
recognised. However, when this allelopathy is viewed in context with the other capability
attributes of camphor laurel the seriousness of the long-term influence of the tree on both
regenerating and established forests can be realised. These factors are now further
summarized.

5.2 The Combined Capability Attributes of Camphor Laurel - An


Increasing Threat to Plant and Animal Biodiversity.
The allelopathy identified as being highly active for camphor laurel in this study (Chapter
2 and 3) combined with the many other capability attributes of the tree (Chapter 1)
indicates that camphor laurel is substantially altering the native vegetation successional
sequence and has potential to increase further its presence, persistence and influence in
the landscape. These attributes which provide for a high stress tolerance as well as
competitive ability in camphor laurel include:
higher leaf areas of the seedlings compared to other plants, canopy dominance
which reduces native plant competition through reduced light levels,
high biomass accumulation which provides for much of the trees stress tolerance,

high longevity enabling persistence in the landscape,

high root competition in the upper soil horizon removing moisture and nutrients
and reducing plant competition,
high competitive ability and rapid growth due to a maximized sugar production
from a biannual leaf renewal at lower altitudes and a highly efficient ability of the
canopy architecture to intercept sunlight,

high biannual seed production and efficient transit over long distances through
varied bird dispersal,
172

rapid growth response following frost, fire or physical damage,


high coppicing ability of the roots, trunk and branches, and a
wide tolerance to various soil conditions (Chapter 1).
Unlike *L. camara, *C. monilifera, *R. fruticosus agg., *P. clandestinum, *C. scoparius
and *A. adenophora identified as being of higher biodiversity threat in New South Wales
than camphor laurel (Chapter 1), this tree has higher biomass accumulation, possesses
persistent canopy dominance to tree height, has a longer recruitment time for seedlings
and saplings, and possesses a much longer life-span. What this means is that camphor
laurel has a longer period of time recruiting in the vegetation before an impact becomes
visually apparent through its dominance as a mature tree, and a greater period of
influence on flora and fauna once mature resulting from a lengthy niche occupation and a
highly dominant canopy. However, a factor that now places camphor laurel with these
species is the presence of allelopathy in the tree which probably all too some degree also
possess (Chapter 1).
Currently, camphor laurel is invading a broad range of vegetation types to a greater
degree than the current vegetation mapping identifies. This is due to the absence of
camphor laurel seedling and sapling mapping studies, which could be used to better
predict threat potential. With mature trees, these are not only seen to be dominating some
plant assemblages but also suppressing communities which require high light levels.
Rainforest plants appear at first to be enhanced by the presence of camphor laurel as
many species including some rare and threatened species germinate and persist below the
dense canopy. These too are identified in this study to be influenced by allelopathy which
may result in an alteration to rainforest succession.
Although camphor laurel appears most often in open habitats the seedlings do have the
potential to germinate and persist as slow growing seedlings under a relatively dense
canopy such as rainforest but are infrequently observed in that vegetation. This is not only
due to the intermediate light-preferring nature of the seedlings, but also likely to the high
predation of seed and seedlings by rats, wallabies and livestock in these less disturbed
forests. However, as time goes by these lowland forests will also undergo invasion as
canopy gaps develop due to tree falls, land slips, fire and anthropogenic influence
173

allowing entry and rapid growth of seedlings from newly arrived seed, as well as light
suppressed seedlings that have survived predation. As the camphor laurel invasion
process is less than 100 years in duration in many areas it is envisaged that over time
many plant communities in the north-eastern New South Wales region will become
dominated by the tree. This study identifies that dominance of the tree will persist through
time placing much stress on some plant species, communities and animals. The tree will
certainly increase its threat status as the invasion process increases over time (Chapter 1,
2 and 4).
In returning to the general hypothesis in Chapter 1 the research has found that:

Camphor laurel exerts a significant effect on the regenerating plant assemblage of


the local region due to its ecology, and

the release of allelopathic compounds from the aqueous leaf leachates of the two
chemo-types, 'cineole' or 'camphor' do contribute to the trees success as an exotic
intrusive element in the flora.

5.3 Management and Further Research


Further research is required to assess the relative roles of allelopathy, plant competition
and stress tolerance in the ecology of camphor laurel as the research concluded that these
aspects allow the tree to be a successful intruder. In other words, which is the more
dominating influence in the success of the tree? Furthermore, trials are needed that
compare the relative importance of these attributes between the camphor and cineole
chemo-types as these where found to vary in their presence in the landscape (Chapter 4).
Is one chemo-type more invasive than the other or are their differences in ecology which
segregate their tolerance to varying environmental conditions? Recruitment studies need
special attention as little work has been undertaken in thickly forested areas to assess
whether the invasion cycle occurring in these closed forest communities represents a long
term threat to the national park system and whether this invasion may influence
productivity in forestry areas. Studies relating to animal interaction such as seed predation,
herbivory and other influences to recruitment would also be useful in understanding how
the process of invasion may be occurring and whether some areas are less prone to
invasion than others. Toxicology studies are also needed to identify if soil biota are
174

influenced and whether the invasion process poses a threat to waterways or particular
organisms.
Of priority is research into improving the management and control of the tree other than
through the use of chemicals as these are expensive for broad scale use and potentially
harmful for human health and the environment depending on the chemical. What is also
needed is an understanding of camphor laurel recruitment in vegetation of less disturbed
character such as National Parks and State Forests and the predictive modelling of the
status of threat to plant communities undergoing invasion.
This research also identifies a few capability attributes of the tree (Chapter 1), which
could be further explored to improve management and control. The seed and seedlings are
highly susceptible to fire and browsing by rats, wallabies and livestock and these limit
recruitment. Such constraints could be useful to consider in management. With fire and
grazing as a method of control for farmers, the burning of leaf litter or intensive grazing
where camphor laurel seedlings are emerging could be useful in assisting in the
prevention of an infestation. In vegetated areas where fire is an integral part of the
ecology, such as dry sclerophyll communities and heath-lands the regular use of a mosaic
burning pattern could also contribute to a broader control of seed and seedlings in some
circumstances. The maintenance of viable wallaby populations in forested areas may also
assist with control as they readily browse seedlings. Shade is also a crucial factor in
seedling development as high shade density reduces the germination and competitive
ability of the seedling. The planting for a fast canopy closure and a substantial groundlayer/mid-layer for light suppression, as well as habitat for camphor laurel seed
destroying wildlife such as the bush rat, various seed grinding birds (Chapter 1, Table 3),
and seedling browsing wallabies may be of assistance in control (Chapter 1).
Another factor which appears to limit the capabilities of the tree is the presence of a
native pathogen which commonly infects mature trees causing partial die-back and even
death in some situations (Plate 1). This infection could be viewed as a constraint on the
growth of the tree (Chapter 1) and investigated as a possible control. The disease has been
observed during this study to occur in both the cineole and camphor chemo-types of
camphor laurel and in moist riparian locations as well as on dry ridge-tops. Previous
work by Firth (1979) identified this disease as Phytopthora sp. but sectioning or culture
175

of the pathogen for positive identification was not attempted. Images of the tree shown in
Plate 1 identify that the infection process begins in the leaves and branch-lets, unlike
Phytopthora. The tree possibly reacts by producing tyloses and gum which then block the
vasculature resulting in the sacrifice of a portion or even the whole tree. It commonly
occurs as a partial die-back followed by epicormic resprouting from the remaining living
branches and trunk (Plate 1c). The process was observed to repeat itself each year on the
infected tree and is referred to here as sequential die-back due to this cyclic nature. In
some situations where damage to the lower trunk of the tree occurs as with the use of
roadside equipment the tree quickly dies, indicating possible infection from soil contact
with the vasculature (Plate 1b). Interestingly, no root infection was observed on any tree
during the study, which would be indicative of Phytopthora infection. The hand
sectioning of leaf material for this research (Chapter 3), may have identified the sexual
fruiting bodies of Rhizopus sp. within the camphor laurel leaf of an infected tree but this
requires further sectioning, culture and taxonomy for positive verification. This same
fungus was later found growing in the top of the extract tanks of both the cineole and
camphor chemo-types prepared in Chapter 2, and then spread to the glasshouse trial
from the extract application where it persisted for several weeks. Further research into
this native disease may be worthy of consideration as a biological control for individual
trees. If such a disease is found, it would reduce the expense of chemical application, be
less hazardous to human health and the environment and allow the removal of an
extensive number of targeted trees. However, further investigation into biological control
agents for the tree in China, Taiwan and Japan is also highly recommended.
Stubbs et al. (1999) identified a large number of economic uses for the camphor laurel
tree which can also be viewed as a method of reducing the trees capacity to influence
flora and fauna. The utilisation of the tree as a further economic resource other than its
current use as timber and a fuel-wood for power generation requires further investigation.
One possible use did emerge from this research that may offer some interest. The
allelochemics of camphor laurel are complex and synergistic in effect and there is
indication in the literature that many of the 38 terpenoids found in the leaves, bark and
wood and the protein cinnamomin found in the seed have insecticidal properties. With
the terpenoids identified in camphor laurel, many have an influence on insect behaviour
and physiology such as being alarm pheromones or substances that interfere with insect
176

(a). Branch-let infection, Uki.

(b). Trunk infection, Uki.

(c). Branch infection with epicormic


resprouting, Casino.

Plate 1. Fungal infection of camphor laurel


at Uki and Casino, NSW of: (a) branch-lets; (b) trunk; (c) branches.

177

healing (Torrsell 1997) or chemical communication (Moritor 2009), and act as


nematicides (Kong et al. 2007). Interestingly the toxic protein cinnamomin which is in
high concentration in the camphor laurel seed has been identified as an efficient
insecticide in studies in China by Zhou et al. (2000), by disrupting the insect central
nervous system. Considering that Australia has a more conducive climate for seed
production with mature trees being able to produce approximately 110,000 seeds (Panetta
2001), this may identify an opportunity for possible use. However, further
experimentation should be undertaken with Australian insect pest species before this
could be considered in addition to the method of seed harvest. Casual observations of the
leaves, bark, soil and leaf litter under the camphor laurel trees visited during this study
has also suggested that invertebrates may occur less frequently and in less diversity than
compared to other canopy dominated vegetation. Apart from possible insecticidal
properties, this also suggests that further research is needed to assess the influence of the
camphor laurel allelochemics on soil and leaf litter organisms as these are an important
food source for many forest animals, and on water quality in slower moving streams or
ponds due to the slow release of terpenoids and phenols from the leaves.
It is also important to consider that removal of large numbers of camphor laurel trees in a
given area for economic or environmental reasons may result in a negative impact on the
ecology as the tree does provide a positive habitat function for many species of flora and
fauna (Chapter 1). Large scale removal would create habitat fragmentation and loss for
some native animal and plant species, soil erosion in some situations and allow the entry
of other weeds including the mass germination of camphor laurel seeds themselves.
Partial and sequential removal with consideration for the existing regeneration, minimal
soil and leaf litter disturbance, plus enrichment planting is recommended to maintain
habitat function when extensively removing camphor laurel forests.

178

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191

Phytochemistry, Allelopathy and the Capability


Attributes of Camphor Laurel (Cinnamomum
camphora (L.) Nees & Eberm.)
in north-eastern New South Wales.

APPENDICES

By John Robert Schenk


B.A. (Hons.) (NSW.)

A thesis submitted for the degree of Doctor of Philosophy:


School of Environmental Science and Management
Southern Cross University

September 2009

LIST OF APPENDICES

A1

Germination of the Trees and Shrubs

A2

Germination of the Vines and Climbers

A3

Germination of the Herbs

A4

Vascular Plant Response Groups

A5

Non-linear Regressions -Trees and Shrubs

A6

Non-linear Regressions Herbs

A7

Non-linear Regressions Vines

A8

Modeled data from the Richards Function

A9

Results of the Deviance Tests

A10

Residual Plots of the Deviance Test

A11

T-test Comparisons of the Glasshouse Data

A12

Summary Effect Tables for the Glasshouse Treatments

A13

Non-linear Regressions Algae

A14

Environmental Site Attributes Field Assessment

A15

T-test Comparisons of the Field Data

ii

Appendix 1: Germination of the Trees and Shrubs.


Group 1: Treatments with higher germination than the control.
Nil
Group 2: One treatment with higher germination than the control,
the other lower.

100
90

% germination

80
70
60
50
40

Control

30

Cineole

20

Cam phor

10
0
1

11

13

15

17

19
Day

21

23

25

27

29

31

33

35

Figure
Figure
A1.1X . G erm ination of E ucalyptus interm edia

10 0

C o ntro l

90

C in e ole

% germination

80

C a m ph or

70
60
50
40
30
20
10
0
1

9 1 0 11 1 2 13 1 4 15 1 6 1 7 18 1 9 20 2 1 22 2 3 24 2 5 26
Day

F ig uA1.2
re X . G erm ination of Lepiderem a pulchella
Figure

100
90

% germination

80
70
60
50

C o n tro l

40

C in e o le

30

Cam phor

20
10
0
1

10

13

16

19

22

25

28

31

34

37

40

43

46

49

52

55

D ay

F i g u A1.3
r e X . G e r m in a t io n o f E u c a ly p t u s s a lig n a
Figure

C o n tro l

90

C in e o le

80

C am phor

% germination

100

70
60
50
40
30
20
10
0
1

11

13

15
17
D ay

19

21

23

25

27

29

31

F ig uA1.4
r e X . G e rm in a tio n o f S y z y g iu m p a n ic u la tu m
Figure

100

C ontrol

90

C ineole

% germination

80

C am phor

70
60
50
40
30
20
10
0
1

11

13

15

17

19
D ay

21

Figure
Figure
A1.5X . G erm ination of Toona australis

A1-2

23

25

27

29

31

33

35

37

Group 3: One treatment with higher germination than the control,


the other with no effect.
100

C ontrol

90

C ineole

80

C am phor

% germination

70
60
50
40
30
20
10
0
1

11

13

15
17
D ay

19

21

23

25

27

29

31

F ig u
re X . G erm in ation o f G u ioa sem iglau ca
Figure
A1.6
100

C ontrol

90

C ineole

% germination

80

C am phor

70
60
50
40
30
20
10
0
1

11

13

15

17

1 9 21
D ay

23

25

27

29

31

33

35

37

F ig uA1.7
re X . G e rm in a tio n o f M a ca ra n g a ta n a riu s
Figure
100

C o n tro l
C in e o le
Cam phor

90

% germination

80
70
60
50
40
30
20
10
0
1

11

21

31

41

51

61

71

81 91 101 111 121 131 141 151 161


D ay

F i gA1.8
u r e X . G e r m in a tio n o f M a llo t u s p h ilip p e n s is
Figure

A1-3

39

100

C o n tro l

90

C in e o le

% germination

80

C am phor

70
60
50
40
30
20
10
0
1

10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64
Day

F ig uA1.9
re XGermination
. G e rm in a tioof
n Ficus
o f F ic u
s m a c ro c a rp a
Figure
macrophylla

Group 4: One treatment with lower germination than the control,


the other with similar levels to the control.
100
90

% germination

80
70
60
50
40

Control
Cineole
Camphor

30
20
10
0
1

7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70
Day

Figure X. Germination of Callistemon viminalis

Figure A1.10 Germination of Callistemon viminalis


100

Control
Cineole
Camphor

90

% germination

80
70
60
50
40
30
20
10
0
1

7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70
Day

Figure
X. Germination
confertus
Figure
A1.11
GerminationofofLophostemon
Lophostemon
confertus

A1-4

Group 5: Treatments with lower germination than the control.


100

C ontrol
C ineole
C am phor

90
80
% germination

70
60
50
40
30
20
10
0
1

11

13

15

17

D ay

19

21

23

25

27

29

31

of Syzygium
S yzygium luehmannii
leuhm an ii
FigA1.12
u re X Germination
. G erm ination of
Figure
100

C o ntro l
C ine ole
C a m p ho r

90

% germination

80
70
60
50
40
30
20
10
0
1

10

13

16

19

22

25

28

31

34

37

40

43

46

49

52

55

Day

m luc id u m
F ig u re X . G erm in a tio n o f L ig u struDay

Figure A1.13 Germination of *Ligustrum lucidum


100

C o n tro l
C in e o le
C am phor

90

% germination

80
70
60
50
40
30
20
10
0
1

11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43
Day

F ig u re X . G e rm in a tio n o f *S o la n u m c a p s ic o id e s

Figure A1.14 Germination of *Solanum capsicoides

A1-5

% germination

100

C o n tro l

90

C in e o le

80

C am phor

70
60
50
40
30
20
10
0
1

11

13

15

17

19

21

23

25

27

29

Day

F ig u
r e X . G e r m in a tio n o f O m a la n th u s p o p u lifo liu s
Figure
A1.15

100

C ontrol
C ineole
C am phor

90

% germination

80
70
60
50
40
30
20
10
0
1

17

25

33

41

49

57

65

73

81 89
D ay

97 105 113 121 129 138 146 154

F igA1.16
u re X . G e rm in ation of H y m e nos poru m flavu m
Figure

1 00

C ontrol
C ineole
C am phor

90

% germination

80
70
60
50
40
30
20
10
0
1

11

21

31

41

51

61

71

81

91 10 1 11 1 121 1 31 1 46 15 6 166 176 1 86


Day

Figure X . G erm ination of *C innam om um cam phora from soil

Figure
Germination
*Cinnamomum
camphora
below aA1.17
C am phor
tree i.e. of
actively
germ inating
seed from soil below a
Camphor laurel tree i.e. actively germinating seed

A1-6

100

Control
Cineole
Camphor

90
80
% germination

70
60
50
40
30
20
10
0
1

11

13

15

18

20

22

26

28

30

34

36

40

42

Day

Figure
X. Germination of Allocasuarina torulosa
Figure
A1.18

Group 6: Treatments with similar level compared to the control


100

Control

90

Cineole

80

Cam phor

% germination

70
60
50
40
30
20
10
0
1

11

13

15
17
D ay

19

21

23

25

27

29

31

FigureFigure
A1.19 X. G erm ination of Cupaniopsis parvifolia
100

C ontrol
C ineole
C am ph or

90

% germination

80
70
60
50
40
30
20
10
0
1

11 13 15 17 19 21 23 2 5 27 29 31 3 3 35 3 7 39 41 43
D ay

F igu
re X . G erm ination of A llocasuarina littoralis
Figure
A1.20

A1-7

100

C ontrol

90

C ineole

% germination

80

C am phor

70
60
50
40
30
20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29
Day

Figure
Figure
A1.21 X . G erm ination of *C assia coluteoides

100

C o n tr o l
C in e o le
C am phor

90

% germination

80
70
60
50
40
30
20
10
0
1

11 14 16 19 21 23 26 28 31 33 35 38 40 42 46 48 50 54 56
Day

F iA1.22
g u r e X . G e r m in a tio n o f A r a u c a r ia c u n n in g h a m ii
Figure

% germination

100

C ontrol

90

C ineole

80

C am phor

70
60
50
40
30
20
10
0
1

11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49
Day

Figure
Figure
A1.23 X . G erm ination of Tristaniopsis laurina

A1-8

Group 7: Treatments fluctuating in effect compared to the control


i.e. a germination delay followed by an increase above the control.

100

C o n tro l
C in e o le
Cam phor

90

% germination

80
70
60
50
40
30
20
10
0
1

7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 76 79
D ay

F ig
u re X . G e rm in a tio n o f F ic u s c o ro n a ta
Figure
A1.24
100

C o n tr o l
C in e o le
C am phor

90

% germination

80
70
60
50
40
30
20
10
0
1

10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64
D ay

F igA1.25
u r e X . G e rm in a tio n o f C o m m e rs o n ia b a r tr a m ia
Figure
100

C o n tro l

90

C in e o le

% germination

80

Cam phor

70
60
50
40
30
20
10
0
1

11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41
D ay

ig u r26
e X . G e r m in a tio n o f A c a c ia m e la n o x y lo n
FigureF A1.

A1-9

100

C ontrol

90

C in eole

% germination

80

C am ph or

70
60
50
40
30
20
10
0
1

9 10 13 14 15 16 17 20 21 22 23 24 28 29 30 31
D ay

FigureFigure
A1.27 X . G erm ination of E ucalyptus m icrocorys

Group 8: Germination below 10 %.


25

Control
Cineole
Camphor

% germination

20
15
10
5
0
1

11

21

31

41

51

61

71
Day

81

91

101

111

121

131

Figure
Figure
A1.28X. Germination of *Lantana camara

25

C ontrol
C ineole
C am phor

% germination

20

15

10

0
1

11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
D ay

ig u re X . G e rm in a tio n o f C o rd ylin e ru b ra
FigureFA1.29

A1-10

25

C o n tro l
C in e o le
C am phor

% germination

20
15
10
5
0
1

11

13

15 17
D ay

19

21

23

25

27

29

31

33

F ig A1.30
u r e X . G e r m in a tio n o f A lp h ito n ia e x c e ls a
Figure

Group 9: Error in control or treatments.


110

Control

100

Cineole - rat predation Rep. 2

90

Camphor

% germination

80
70
60

Rep.1 only

50
40
30
20
10
0
1

11

13

15

17 19
Day

21

23

25

27

29

31

33

FigureFigure
A1.31X. Germination of Davidsonia pruriens

100

Control - N & P excess

90

Cineole

% germination

80

Camphor

70
60
50
40
30
20
10
0
1

13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85 89 93
Day

Figure
Melia
azedarach
GerminationofofMelia
Figure
A1.32X. Germination
azedarach

A1-11

100

Control - seed dessication


Cineole
Camphor

90
80
% germination

70
60
50
40
30
20
10
0
1

17 25 33 41 49 57 65 73 81 89 97 105 113 121 129 137 145 153 163 171


Day

Figure.
Figure
A1.33 X Germination of *Cinnamomum camphora from
depulped seed of fallen ripe fruit.
100

C ontrol - N & P e xce ss

90

C ineole

% germination

80

C am phor

70
60
50
40
30
20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51 53
D ay

ig u re X . G e rm ina tio n o f F lind e rsia xa nth o xyla


FigureFA1.34

100

Control - N & P excess

% germination

90
80

Cineole

70

Camphor

60
50
40
30
20
10
0
1

9 11 13 15 17 21 23 25 30 32 36 38 43 46 50 53 56 58 62
Day

FigureFigure
A1.35 X. Germination of Castanospermum australe

A1-12

100

Control - N & P excess

90

Cineole

80
Camphor

% germination

70
60
50
40
30
20
10
0
1

10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73
Day

FigureFigure
A1.36 X. Germination of Jagera pseudorhus

100

C ontrol - waterlogging

90

C ineole

80

C am phor

% germination

70
60
50
40
30
20
10
0
1

11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49
D ay

FigureFigure
A1.37 X. G erm ination of D iploglotis australis

A1-13

Appendix 2: Germination of the Vines and Climbers.


Group 1: Treatments with higher germination than the control.
100

C o n tr o l
C in e o le
Cam phor

90

% germination

80
70
60
50
40
30
20
10
0
1

F ig A2.1
u re X .
Figure

10 11 12 13 14 15 16 17 18 19 20 21 22
Day

G e rm in a tio n o f * C a rd io s p e rm u m g ra n d iflo ru m

Group 2: One treatment with higher germination than the control,


the other lower.
Nil

Group 3: One treatment with higher germination than the control,


the other similar to the control.
Nil

Group 4: One treatment with lower germination than the control,


the other similar to the control.
Nil

Group 5: Treatments with lower germination than the control.

100

Control
Cineole
Camphor

90
80
% germination

70
60
50
40
30
20
10
0
1

9 13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85 89 93 97
Day

Figure
Figure
A2.2X. Germination of *Anredera cordifolia

Group 6: Treatments with effect similar to the control.


Nil

Group 7: Treatment(s) fluctuating in effect compared to the control


i.e. a germination delay followed by an increase above the control.

100

Control
Cineole
Camphor

90

% germination

80
70
60
50
40
30
20
10
0
1

13

19

25

31

37

43

49

55

61

67

73

Day

Figure
Figure
A2.3X. Germination of Cissus hypoglauca

A2-2

79

85

91

97 103 109 115 121

Group 8: Germination below 10 %


100

C ontrol
C ineole
C am phor

90

% germination

80
70
60
50
40
30
20
10
0
1

15

22

29

36

43

50

57

64

71

78

85

92

99 106 113 120 127 134 141

D ay

Figure
Figure
A2.4X. G erm ination of G eitonoplesium cym osum

100

C o n tro l
C in e o le
Cam phor

90

% germination

80
70
60
50
40
30
20
10
0
1

7 1 0 1 3 1 6 1 9 2 2 2 5 2 8 3 1 3 4 3 7 4 0 4 3 4 6 4 9 5 2 5 5 5 8 6 1 64 6 7 7 0 7 3
Day

F ig
u re X . G e rm in a tio n o f *P a s s iflo ra s u b e ro s a
Figure
A2.5

Group 9: Error in control or treatments.


Nil

A2-3

Appendix 3: Germination of the Herbs.


Group 1: Treatments with higher germination than the control.
100

Control

90

Cineole
Camphor

% germination

80
70
60
50
40
30
20
10
0
1

10

13

16

19

22

25

28

31

34

37

40

43

46

49

52

55

58

61

Day

Figure
A3.1X. Germination of Oplismenus aemulus
Figure

Group 2: One treatment with higher germination than the control,


the other lower.
Nil

Group 3: One treatment with higher germination than the control,


the other similar to the control.
1 00

C on tro l

90

C ine o le

80

C am ph or

% germination

70
60
50
40
30
20
10
0
1

15

22

29

36

43

50

57

64

71

78

85

92

D ay

F ig uA3.2
re X . G e rm in a tio n o f C yp e ru s e n e rv is
Figure

9 9 1 0 6 1 13 1 20 12 7 13 4

100

C o n tro l
C in e o le
C am phor

90

% germination

80
70
60
50
40
30
20
10
0
1

11

13

15

17
D ay

19

21

23

25

27

29

31

33

F ig u
re X . G e rm in a tio n o f *S e ta ria s p h a c e la ta v a r. n a ro k
Figure
A3.3

100

C o n tro l

90

C in e o le

% germination

80

C am phor

70
60
50
40
30
20
10
0
1

10

13

16

19

22

25

28

31

34

37

40

43

46

49

52

55

58

Day

F ig uA3.4
re X . G e rm in a tio n o f O tto c h lo a g ra c illim a
Figure

Group 4: One treatment with lower germination than the control,


the other similar to the control.
100

C o n tro l
C in e o le
Cam phor

90

% germination

80
70
60
50
40
30
20
10
0
1

17

25

33

41

49

57

65 73
Day

81

89

F ig uA3.5
r e X . G e rm in a tio n o f A lp in ia c a e ru le a
Figure

A3-2

97

105 113 121 129 137

100

C o n tr o l
C in e o le
C am phor

90
80
% germination

70
60
50
40
30
20
10
0
1

11

13

15

17

19

21

23

25

D ay

F i g uA3.6
r e X . G e r m in a t io n o f * C h lo r is g a y a n a
Figure

Group 5: Treatments with lower germination than the control, with


some returning to near control levels.
100

C o n tro l
C in e o le
C am phor

90

% cutting strike

80
70
60
50
40
30
20
10
0
1

11

13

15

17
D ay

19

21

23

25

27

29

31

F i g u r e X . S tr ik e o f V io la h e d e r a c e a

Figure A3.7 Strike of Viola hederaceae

100
90

% germination

80
70
60
50
40
30

Control
Cineole
Cam phor

20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25
Day

Figure
X. Germination of *Pennisetum clandestinum
Figure
A3.8

A3-3

100

C o n tr o l
C in e o le
Cam phor

90
80
% cutting strike

70
60
50
40
30
20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43
Day

F ig A3.9
u r e X . S tr ik e o f P ra tia p u rp u re s c e n s
Figure

100

C o n tro l
C i n e o le
Cam phor

90

% germination

80
70
60
50
40
30
20
10
0
1

13

17

21

25

29

33 37
D ay

41

45

49

53

57

61

65

69

F i g A3.10
u r e X . G e r m i n a t i o n o f * P a s p a lu m d il a t a t u m
Figure

100

Control
Cineole
Cam phor

90

% germination

80
70
60
50
40
30
20
10
0
1

7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67
Day

Figure
Figure
A3.11X. Germination of Helychrysum bracteatum

A3-4

Group 6: Treatments with similar effect compared to the control.


Nil

Group 7: Treatments fluctuating in effect compared to the control


i.e. a germination delay followed by an increase above the control.
100

C o n tr o l
C in e o le
C am phor

90
80
% germination

70
60
50
40
30
20
10
0
1

15

22

29

36

43

50

57
D ay

64

71

78

85

92

F ig A3.12
u r e . X G e rm in a tio n o f L o m a n d ra lo n g ifo lia
Figure

Group 8: Germination below 10 %


Nil

Group 9: Error in control or treatments.


Nil

A3-5

99

105 112

% of species in life-form group

Appendix 4: Vascular Plant Response Groups.


50
45
40
35
30
25
20
15
10
5
0

vines/twinners (species n = 7)

43%

herbs (species n = 19)


trees/shrubs (species n =43)
29%
21%

28%
21%

21%

14 %

14%

16%

11%
7%

16%

7%

5%

5%

0%

0%

Group 1:
Stim/Return

16%

14%

12%

Group 2:
Stim/Delay

Group 3:
Full Stim

0%

Group 4:
Delay/Return

Group 5:
Delay/Stim

Group 6:
Full Delay

Group 7:
Similar to Control

Response group

Response group

Weight of 100 propagules (grms)

Weight of 100 propagules (g)

Figure X. Proportion of species represented as life-forms in


Figure A4.1 Proportion of species represented as life-forms in each
each response groups.
response group.
60

seeds
vegetative propagules

50

n = 44

40
30
20
10
0
Group 1:
Stim/Return

Group 2:
Stim/Delay

Group 3:
Full Stim

Group 4:
Delay/Return

Group 5:
Delay/Stim

Group 6:
Full Delay

Response group

Group 7:
Similar to
Control

% of species in propagule group

FigureFigure
A4.2 Propagule
weights
in theinresponse
groups.groups.
weights
the response
X. Propagule

35
30
25

32%

dry propagules (species n = 42)


moist propagules (sample n =
28)

24%

18%

20

17%
14%

15
10

14%
11%

10%

14%

14%
11%

7% 7%

7%

5
0
Group 1:
Stim/Return

Group 2:
Stim/Delay

Group 3:
Full Stim

Group 4:
Delay/Return

Response group

Group 5:
Delay/Stim

Group 6:
Full Delay

Group 7:
Similar to
Control

Figure X. Proportion of dry and moist propagules in each


Figure A4.3 Proportion of dry and moist propagules in each response group.
response group.

% of species in origin group

35

Introduced plants (species n =17)

30

29%

Native plants (species n = 52)

25

27%

23%

20

18%
15%

15

12%

15%
12% 12%

11%

10%

10%

10

6%

5
0%

0
Group 1:
Stim/Return

Group 2:
Stim/Delay

Group 3:
Full Stim

Group 4:
Delay/Return

Response group

Group 5:
Delay/Stim

Group 6:
Full Delay

Group 7:
Similar to Control

FigureProportion
X. Proportion
of introduced
and
nativeplants
plantsinineach
each
Figure A4.4
of introduced
and
native
response
responsegroup.
group.

% of species in similarity
response group

30

28%

similar response to 'cineole' and


'camphor' leachate (species n = 19)
distinct response (species n = 50)

25

26%

26%

20
16%

16%

16% 16%

16%

15

12%

11%

10
6%

6%

5%

5
0%

0
Group 1:
Stim/Return

Group 2:
Stim/Delay

Group 3:
Full Stim

Group 4:
Delay/Return

Group 5:
Delay/Stim

Group 6:
Full Delay

Group 7:
Similar to Control

Response group

% of species in germination
group

X. Proportion
of species
with
similarand
anddistinct
distinctresponses to
Figure Figure
A4.5 Proportion
of species
with
similar
responses to the
two
leachates
in
each
response
the two leachates in each responsegroup.
group.

45
40
35
30
25
20
15
10
5
0

species displaying dormancy (species n = 20)


rapidly germinating species (species n = 49)

40%

25%
20.50%

20.50%

10%

12%

16.50%

14.50%
10%

10%
6%

10%
5%
0%

Group 1:
Stim/Return

Group 2:
Stim/Delay

Group 3:
Full Stim

Group 4:
Delay/Return

Group 5:
Delay/Stim

Group 6:
Full Delay

Response group

Figure
A4.6X.Proportion
each response group
groupwith
with
Figure
Proportionof
of species
species in each
dormancy
and
rapid
germination.
dormancy and rapid germination.

A4-2

Group 7:
Similar to Control

Appendix 5. Non-Linear Regressions - Trees and Shrubs


using the Richards function.
100
90

% germination

80
70
60
50
40

r = 0.9868

30
20

observed germination

10

fitted germination

0
1

11

13

15

17

19

21

23

25

27

29

31

33

Days

FigureFigure
A5.1a1a. Comparison of actual and fitted germination for the 'control'
treatment of Acacia melanoxylon .
100
90
% germination

80
70
60
50
2

r = 0.965227

40
30
20

observed germination
fitted germination

10
0
1

11

13

15

17

19

21

23

25

27

29

31

33

Days

FigureFigure
A5.1b
1b. Comparison of actual and fitted germination for the 'cineole'
treatment of Acacia melanoxylon.
100
90

% germination

80
70
60

r = 0.991812

50
40
30
20

observed germination

10

fitted germination

0
1

11

13

15

17

19

21

Days

23

25

Figure A5.1c
Figure 1c. Comparison of actual and fitted germination for the
'camphor' treatment of Acacia melanoxylon.

27

29

31

33

100
90
% germination

80
70
60
50
40
30

r2 = 0.933157

20

observed germination

10

fitted germination

0
1

11

13

15

17

19

21

23

25

27

29

31

33

Days

FigureFigure
A5.2a
2a. Comparison of actual and fitted germination for the 'control'
treatment of Allocasuarina littoralis.

100
90

% germination

80
70
60
50
40
2

r = 0.975398

30
20

observed germination

10

fitted germination

0
1

11

13

15

17

19

21

23

25

27

29

31

33

Days

2b. Comparison of actual and fitted germination for the 'cineole'


FigureFigure
A5.2b

treatment of Allocasuarina littoralis.

100
90

% germination

80
70
60
50
40

r2 = 0.970501

30
20

observed germination
fitted germination

10
0
1

11

13

15

17

19

21

23

25

FigureFigure
A5.2c2c. Comparison of actual and fittedDays
germination for the
'camphor' treatment of Allocasuarina littoralis.

A5-2

27

29

31

33

100
observed germination

90

fitted germination

% germination

80
70
60
50
40

r = 0.990662

30
20
10
0
1

11

13

15

17

19

21

23

25

27

29

31

33

Days

FigureFigure
A5.3a3a. Comparison of actual and fitted germination for the 'control'
treatment of Allocasuarina torulosa.

100
observed germination
fitted germination

90
% germination

80
70
60
50
40
30

r2 = 0.938912

20
10
0
1

11

13

15

17

19

21

23

25

27

29

31

33

29

31

33

Days

3b. Comparison of actual and fitted germination for the 'cineole'


FigureFigure
A5.3b

treatment of Allocasuarina torulosa.

100

observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.971712

20
10
0
1

11

13

15

17

19

21

Days

23

25

Figure A5.3c
Figure 3c. Comparison of actual and fitted germination for the
'camphor' treatment of Allocasuarina torulosa.

A5-3

27

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.988006

20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days

4a. Comparison of actual and fitted germination for the 'control'


FigureFigure
A5.4a

treatment of Araucaria cunninghamii.

100

observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.987405

20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days

4b. Comparison of actual and fitted germination for the 'cineole'


FigureFigure
A5.4b

treatment of Araucaria cunninghamii.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r2 = 0.992117

10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days

Figure A5.4c
Figure 4c. Comparison of actual and fitted germination for the
'camphor' treatment of Araucaria cunninghamii.

A5-4

100
90

% germination

80
70

r2 = 0.977949

60
50
40
30
20

observed germination

10

fitted germination

0
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51
Days

5a. Comparison of actual and fitted germination for the 'control'


FigureFigure
A5.5a

treatment of Callistemon viminalis.

100
90

% germination

80
70
60
50
40

r2 = 0.992851

30
20
observed germination

10

fitted germination

0
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51
Days

Figure Figure
A5.5b5b. Comparison of actual and fitted germination for the 'cineole'
treatment of Callistemon viminalis.

100
90

% germination

80
70
60

r2 = 0.962731

50
40
30
20

observed germination

10

fitted germination

Figure

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51
Days
A5.5c
Figure 5c. Comparison of actual and fitted germination for the
'camphor' treatment of Callistemon viminalis.

A5-5

100
observed germination
fitted germination

90

% germination

80
70
60
50
40
30

r2 = 0.997102

20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
Days

6a. Comparison of actual and fitted germination for the 'control'


FigureFigure
A5.6a

treatment of *Cassia coluteoides.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30

0.999402
r2 = 1.000598

20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
Days

6b. Comparison of actual and fitted germination for the 'cineole'


FigureFigure
A5.6b

treatment of *Cassia coluteoides.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.998606

20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
Days

Figure A5.6c
Figure 6c. Comparison of actual and fitted germination for the
'camphor' treatment of *Cassia coluteoides.

A5-6

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r = 0.992159

10
0
1

15 22 29 36 43 50 57 64 71 78 85 92 99 106 113 120 127 134


Days

Figure 7a. Comparison of actual and fitted germination for the 'control'
Figure A5.7a
Comparison
of actual
and fitted
for
camphora
treatment
of Cinnamomum
fromgermination
soil.

the
control treatment of *Cinnamomum camphora from soil.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
2

30

r = 0.998766

20
10
0
1

15 22 29 36 43 50 57 64 71 78 85 92 99 106 113 120 127 134


Days

Figure 7b. Comparison of actual and fitted germination for the 'cineole'
Comparison of actual and fitted germination for the
Figure A5.7b
treatment of Cinnamomum camphora from soil.

cineole treatment of *Cinnamomum camphora from soil.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.937535

20
10
0
1

15 22 29 36 43 50 57 64 71 78 85 92 99 106 113 120 127 134


Days

Figure 7c. Comparison od actual and fitted germination for the


Comparison
of actualcamphora
and fittedfrom
germination
for the camphor
Figure
A5.7c
'camphor'
treatment
of Cinnamomum
soil.

treatment of *Cinnamomum camphora from soil.

A5-7

100
90

% germination

80
70

r2 = 0.985176

60
50
40
30
20

observed germination
fitted germination

10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51
Days

FigureFigure
A5.8a
8a. Comparison of actual and fitted germination for the 'control'
treatment of Commersonia bartramia.

100
90
% germination

80
70

r2 = 0.987748

60
50
40
30
20

observed germination

10

fitted germination

0
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51
Days

8b. Comparison of actual and fitted germination for the 'cineole'


FigureFigure
A5.8b

treatment of Commersonia bartramia.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r2 = 0.976206

10
0

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51

Figure A5.8c

Days

Figure 8c. Comparison of actual and fitted germination for the


'camphor' treatment of Commersonia bartramia.

A5-8

100
90
2

% germination

80

r = 0.995028

70
60
50
40
30
20

observed germination

10

fitted germination

0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25

Days

9a. Comparison of actual and fitted germination for the 'control'


FigureFigure
A5.9a

treatment of Cupaniopsis parvifolia.

100
90
% germination

80

0.995319
r = 1.004681

70
60
50
40
30
20

observed germination

10

fitted germination

0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25

Days

FigureFigure
A5.9b9b. Comparison of actual and fitted germination for the 'cineole'
treatment of Cupaniopsis parvifolia .

100
90

% germination

80
70

r2 = 0.991319

60
50
40
30
20

observed germination

10

fitted germination

0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25
Days

Figure A5.9c
Figure 9c. Comparison of actual and fitted germination for the
'camphor' treatment of Cupaniopsis parvifolia.

A5-9

100
90

% Germination

80
70

r2 = 0.99234

60
50
40
30
20

observed germination

10

fitted germination

0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27

Days

10a. Comparison of actual and fitted germination for the


Comparison of actual and fitted germination for the
FigureFigure
A5.10a
'control' treatment of Eucalyptus intermedia.

control treatment of Eucalyptus intermedia.

100
90

% Germination

80
70

r2 = 0.97097

60
50
40
30

observed germination
fitted germination

20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27

Days
Days
Figure 10b. Comparison of actual and fitted germination for the
Figure A5.10b'cineole'
Comparison
of
actual
and
fitted
germination for the
treatment of Eucalyptus
intermedia.

% Germination

cineole treatment of Eucalyptus intermedia.

100
90
80
70
60
50
40
30
20
10
0

r2 = 0.98914

observed germination
fitted germination

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27
Days

Comparison of actual and fitted germination for the


FigureFigure
A5.10c
10c. Comparison of actual and fitted germination for the
camphor treatment of Eucalyptus intermedia
'camphor' treatment of Eucalyptus intermedia.

A5-10

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r = 0.994543

10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
Days

Figure 11a. Comparison of actual and fitted germination for the


Figure A5.11a

'control' treatment of Eucalyptus microcorys.

100

observed germination
fitted germination

90

% germination

80
70
60
50
40
30
2

20

r = 0.977238

10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
Days

Figure A5.11b
Figure 11b. Comparison of actual and fitted germination for the
'cineole' treatment of Eucalyptus microcorys.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r2 = 0.994289

10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
Days

Figure

Figure 11c. Comparison of actual and fitted germination for the


A5.11c 'camphor''
Comparison
of actual
and fitted germination
microcorys. for the camphor
treatment
of Eucalyptus

treatment of Eucalyptus microcorys

A5-11

100
90
% germination

80
70

r2 = 0.969317

60
50
40
30
20

observed germination

10

fitted germination

0
1

11

13

15

17

19

21 23
Days

25

27

29

31

33

35

37

39

41

FigureFigure
A5.12a
12a. Comparison of actual and fitted germination for the
'control' of Eucalyptus saligna .

100
90

% germination

80
70

r2 = 0.979079

60
50
40
30
20

observed germination

10

fitted germination

0
1

11 13

15

17

19

Figure A5.12b

21

23

Days

25

27

29

31

33

35

37

39

41

Figure 12b. Comparison of actual and fitted germination for the


'cineole' treatment of Eucalyptus saligna.

100
90

r2 = 0.963965

% germination

80
70
60
50
40
30
20

Observed Germination

10

Fitted Germination

0
1

11

13

15

17 19

21

23

25

27

29

31

33

Days
FigureFigure
A5.12c
12c. Comparison of actual and fitted germination for the
'camphor' treatment of Eucalyptus saligna.

A5-12

35

37

39

41

100
observed germination
fitted germination

90
% germination

80
70
60

r = 0.986813

50
40
30
20
10
0
1

10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61
Days

Figure 13a. Comparison of actual and fitted germination for the


Figure A5.13a

'control' treatment of Ficus coronata.

r2 = 0.997163
100
90

% germination

80
70
60
50
40
30
20

observed germination

10

fitted germination

0
1

10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61
Days

Figure 13b. Comparison of actual and fitted germination for the


Figure A5.13b

'cineole' treatment of Ficus coronata.

100
90

% germination

80

0.995592
r = 1.004308

70
60
50
40
30
20

observed germination

10

fitted germination

0
1

10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61
Days

Figure 13c. Comparison of actual and fitted germination of the


Figure A5.13c

'camphor' treatment of Ficus coronata.

A5-13

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r = 0.961714

10
0
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51 53
Days

14a. Comparison of actual and fitted germination for the 'control'


FigureFigure
A5.14a

treatment
treatmentofofFicus
Ficusmacrocarpa.
macrophylla.

100

observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.957777

20
10
0

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51 53
Days

Figure 14b. Comparison of actual and fitted germination for the


Figure A5.14b

macrocarpa .
'cineole'
treatment
of Ficus
treatment
of Ficus
macrophylla.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r = 0.964281

10
0

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51 53
Days

Figure 14c. Comparison of actual and fitted germination for the


Figure A5.14c

'camphor'
treatment of Ficus macrocarpa .
camphor treatment of Ficus macrophylla.

A5-14

100
90

% germination

80
70
60

r2 = 0.993688

50
40
30
20

observed germination

10

fitted germination

0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
Days

Figure 15a. Comparison of actual and fitted germination for the


Figure A5.15a

'control' treatment of Guioa semiglauca.

100
90

% germination

80
70
2

r = 0.993134

60
50
40
30
20

observed germination

10

fitted germination

0
1

10 11 12 13 14 15 16 17 18 19 20 21 22 23 24

Days

Figure 15b. Comparison of actual and fitted germination for the


Figure A5.15b

'cineole' treatment of Guioa semiglauca .

100
90

% germination

80

r = 0.991381

70
60
50
40
30
20

observed germination

10

fitted germination

0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
Days

Figure A5.15c
Figure 15c. Comparison of actual and fitted germination for the
'camphor' treatment of Guioa semiglauca.

A5-15

100
90

% germination

80
70
60

r2 = 0.999709

50
40
30
20

observed germination

10

fitted germination

13

19

25

31

37

43

49

55

61

67

73

79

85

91

97 103 109 115

Days

Figure 16a. Comparison of actual and fitted germination for the


Figure A5.16a

'control' treatment of Hymenosporum flavum.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r = 0.985642

10
0
1

13 19 25 31 37 43 49 55 61 67 73 79 85 91 97 103 109 115


Days

Figure A5.16b
Figure 16b. Comparison of actual and fitted germination for the
'cineole' treatment of Hymenosporum flavum.

100

observed germination

90

fitted germination

% germination

80
70
60
50
40

r2 = 0.989331

30
20
10
0
1

13 19 25 31 37 43 49 55 61 67 73 79 85 91 97 103 109 115


Days

Figure A5.16c
Figure 16c. Comparison of actual and fitted germination for the
'camphor' treatment of Hymenosporum flavum.

A5-16

100

observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.993764

20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
Days

Figure A5.17a
Figure 17a. Comparison of actual and fitted germination for the
'control' treatment of *Ligustrum lucidum.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40

r2 = 0.997358

30
20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23

Days

Figure A5.17b
Figure 17b. Comparison of actual and fitted germination for the
'cineole' treatment of *Ligustrum lucidum.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40

r2 = 0.999097

30
20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
Days

Figure A5.17c
Figure 17c. Comparison of actual and fitted germination for the
'camphor' treatment of *Ligustrum lucidum.

A5-17

100
90

% germination

80
70

r2 = 0.9933

60
50
40
30
20

observed germination

10

fitted germination

0
1

10

11

12

13

14

15

16

17

18

Days

Figure 18a. Comparison of actual and fitted germination for the


Figure A5.18a
Comparison of actual and fitted germination for the control
'Control' treatment of Lepiderema pulchella
treatment of Lepiderema pulchella.

100
90

% germination

80
70

r2 = 0.9933

60
50
40
30
20

observed germination
fitted germination

10
0
1

10

11

12

13

14

15

16

17

18

Days

18b. Comparison of actual and fitted germination for the


Figure Figure
A5.18b
Comparison of actual and fitted germination for the cineole
'Cineole' treatment of Lepiderema pulchella
treatment of Lepiderema pulchella.

100
90

% germination

80

r2= 0.999713

70
60
50
40
30
20

observed germination
fitted germination

10
0
1

10

11

12

13

14

15

16

Days

Figure 18c. Comparison of actual and fitted germination for the


Figure A5.18c
Comparison of actual and fitted germination for the camphor
'camphor' treatment of Lepiderema pulchella .
treatment of Lepiderema pulchella.

A5-18

17

18

100

observed germination

90

fitted germination

80
% germination

70
60
50
40
30

r = 0.975612

20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days

Figure 19a. Comparison of actual and fitted germination for the


Figure A5.19a

'control' treatment of Lophostemon confertus.

100
observed germination

90

fitted germination

80
% germination

70
60
50
40
30

r = 0.910401

20
10
0
1

Figure

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days
A5.19b
Figure 19b. Comparison of actual and fitted germination for the
'cineole' treatment' of Lophostemon confertus .

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
2

r = 0.91715

30
20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days
FigureFigure
A5.19c
19c. Comparison of actual andfitted germination for the
'camphor' treatment of Lophostemon confertus .

A5-19

100
90

% germination

80
70
60
50

r = 0.9987

40
30
20

observed germination

10

fitted germination

0
1

11

13

15

17 19
Days

21

23

25

27

29

31

33

Figure A5.20a
Figure 20a. Comparison of actual and fitted germination fot the
'control' treatment of Macaranga tanarius .

100
90

% germination

80
70

r2 = 0.995201

60
50
40
30
20

observed germination

10

fitted germination

0
1

11

13

15

17

19

21

23

25

27

29

31

33

Days

Figure A5.20b
Figure 20b. Comparison of actual and fitted germination for the
'cineole' treatment of Macaranga tanarius.

100
90

% germination

80
70
60

r2 = 0.990722

50
40
30
20

observed germination

10

fitted germination

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33
Days

Figure A5.20c
Figure 20c. Comparison of actual and fitted germination for the
'camphor' treatment of Macaranga tanarius.

A5-20

100
observed germination

90

fitted germination

80
% germination

70
60
50
40
30
20

r2 = 0.974462

10
0
1

9 13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85 89 93
Days
A5.21a
Figure 21a. Comparison of actual and fitted germination for the 'control'
treatment of Mallotus philippensis.

Figure

100

observed germination

90

fitted germination

% germination

80
70
60
50
40

r2 = 0.977562

30
20
10
0
1

9 13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85 89 93
Days

Figure 21b. Comparison of actual and fitted germination for the 'cineole'
Figure A5.21b
treatment of Mallotus philippensis.

% germination

100
90

observed germination

80

fitted germination

70
60
50
40
30

r2 = 0.970684

20
10
0

1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 76 79 82 85 88 91
Days

Figure Figure
A5.21c
21c. Comparison

of actual and fitted germination for the


'camphor' treatment of Mallotus phillippensis.

A5-21

100
90
% germination

80
70

r2 = 0.99441

60
50
40
30
20

observed germination
fitted germination

10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25
Days

Figure 22a. Comparison of actual and fitted germination for the


Figure A5.22a

'control' treatment of Omalanthes populifolius.

100
90

% germination

80
70
60

r = 0.982691

50
40
30
20

observed germination

10

fitted germination

0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25
Days

Figure 22b. Comparison of actual and fitted germination for the


Figure A5.22b

'cineole' treatment of Omalanthes populifolius.

100
90

% germination

80
70

r2 = 0.997796

60
50
40
30
20

observed germination

10

fitted germination

0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25
Days

Figure A5.21c
Figure 22c. Comparison of actual and fitted germination for the
'camphor' treatment of Omalanthes populifolius.

A5-22

100
90

% germination

80
2

70

r = 0.997965

60
50
40
30
20

observed germination
fitted germination

10
0
1

11

13

15

17

19

21

23

25

27

29

31

33

Days

Figure A5.23a
Figure 23a. Comparison of actual and fitted germination for the
'control' treatment of Solanum capsicoides.

80
70
% germination

60

r2 = 0.995957

50
40
30
20

observed germination
fitted germination

10
0
1

11

13

15

17 19
Days

21

23

25

27

29

31

33

Figure A5.23b
Figure 23b. Comparison of actual and fitted germination for the
'cineole' treatment of Solanum capsicoides.

100
90

% germination

80
70
60
2

50

r = 0.980759

40
30
20
observed germination

10

fitted germination

0
1

11

13

15

17

19

21

23

25

Days

FigureFigure
A5.23c
23c. Comparison of actualand fitted germination for the
'camphor' treatment of Solanum capsicoides.

A5-23

27

29

31

33

100
90
2

r = 0.995336

% germination

80
70
60
50
40
30
20

observed germination

10

fitted germination

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26
Days

Figure 24a. Comparison of actual and fitted germination of the 'control'


Figure A5.24a
Comparison of actual and fitted germination for the control

treatment
of Syzygium
leuhmanii.
treatment
of Syzygium
luehmannii..

100
90

% germination

80

r = 0.994057

70
60
50
40
30
20

observed germination

10

fitted germination

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26
Days

Figure 24b. Comparison of actual and fitted germination for the


Comparison
of actual
and of
fitted
germination
for the control
'cineole'
treatment
Syzygium
leuhmanii.
treatment of Syzygium luehmannii.

Figure A5.24b

100
90

r2 = 0.987556

% germination

80
70
60
50
40
30
20

observed germination

10

fitted germination

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26
Days

Figure 24c. Comparison of actual and fitted germinination for the


Figure A5.24c
Comparison of actual and fitted germination for the control

'camphor'
treatment luehmannii..
of Syzygium leuhmanii.
treatment
of Syzygium

A5-24

100
90

% germination

80

r2 = 0.982773

70
60
50
40
30
20

observed germination
fitted germination

10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27

Days

Figure Figure
A5.25a25a. Comparison of actual and fitted germination for the
'control' treatment of Syzygium paniculatum .

100
90

% germination

80

r2 = 0.989232

70
60
50
40
30
20

observed germination

10

fitted germination

0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26

Days

Figure A5.25b
Figure 25b. Comparison of actual and fitted germination for the
'cineole' treatment of Syzygium paniculatum .

100
90

r2 = 0.975956

% germination

80
70
60
50
40
30
20

observed germination

10

fitted germination

0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26

Days

Figure A5.25c
Figure 25c. Comparison of actual and fitted germination for the
'camphor' treatment of Syzygium paniculatum.

A5-25

100
90

% germination

80

r2 = 0.984527

70
60
50
40
30
20

observed germination

10

fitted germination

0
Days

Figure A5.26a
Figure 26a. Comparison of actual and fitted germination for the
'control' treatment of Toona australis .

100
90

% germination

80

r2 = 0.994781

70
60
50
40
30
20

observed germination

10

fitted germination

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31

Days

Figure A5.26b
Figure 26b. Comparison of actual and fitted germination for
the'cineole' treatment of Toona australis .

100
90

% germination

80
70

r2 = 0.99564

60
50
40
30
20

observed germination
fitted germination

10
0

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31

Days
Figure 26c. Comparison of actual and fitted germination for the 'camphor'
Figure A5.26c
treatment of Toona australis.

A5-26

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
2

30

r = 0.965143

20
10
0
1

11

13

15

17

19

21

23

25

27

29

31

33

35

Days

Figure A5.27a
Figure 27a. Comparison of actual and fitted germination for the
'control' treatment of Tristaniopsis laurina.
100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.982399

20
10
0
1

11

13

15

17

19

21

23

25

27

29

31

33

35

Days

Figure A5.27b
Figure 27b. Comparison of actual and fitted germination for the
'cineole' treatment of Tristaniopsis laurina.
100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.990153

20
10
0
1

11

13

15

17

19

21

23

25

27

29

31

33

35

Days

Figure A5.27c
Figure 27c. Comparison of actual and fitted germination for the
'camphor' treatment of Tristaniopsis laurina.

Note: *Lantana camara is not included due to low germination of 2% in the control,
0% cineole treatment and 6% camphor treatment.

A5-27

Appendix 6.

Non-Linear Regressions - Herbs


using the Richards function.

100

observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r = 0.991167

20
10
0
1

10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61
Days

Figure 1a. Comparison of actual and fitted germination for the 'control'
Figure A6.1a

treatment of Alpinea caerulea.


100
observed germination
fitted germination

90

% germination

80
70
60
50
40

r2 = 0.97487

30
20
10
0
1

10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61
Days

Figure Figure
A6.1b1b. Comparison of actual and fitted germination for the 'cineole'
treatment of Alpinea caerulea.
100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.974085

20
10
0
1

10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61
Days

FigureFigure
A6.1c1c. Comparison of actual and fitted germination for the
'camphor' treatment of Alpinea caerulea.

100
observed germination
fitted germination

90

% germination

80
70
60
50
40
30
20

r2 = 0.991274

10
0
1

10 11 12 13 14 15 16 17 18 19 20
Days

Figure Figure
A6.2a2a. Comparison of actual and fited germination for the 'control'
treatment of *Chloris gayana.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r2 = 0.966286

10
0
1

10 11 12 13 14 15 16 17 18 19 20
Days

Figure 2b. Comparison of actual anf fitted germination for the 'cineole'
Figure A6.2b

treatment of *Chloris gayana.

100
observed germination
fitted germination

90

% germination

80
70
60
50
40
30
20

r2 = 0.865072

10
0
1

10 11 12 13 14 15 16 17 18 19 20
Days

Figure Figure
A6.2c2c. Comparison of actual and fitted germination for the
'camphor' treatment of *Chloris gayana.

A6-2

100
90

% germination

80

observed germination
fitted germination

70
60
50
40
30

r2 = 0.998258

20
10
0
1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 76 79 82 85
Days

Figure Figure
A6.3a3a. Comparison of actual and fitted germination for the 'control'
treatment of Cyperus enervis.

100
90

% germination

80

observed germination
fitted germination

70
60
50
40
30

r2 = 0.949762

20
10
0
1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 76 79 82 85
Days

Figure 3b. Comparison of actual and fitted germination for the 'cineole'
Figure A6.3b

treatment of Cyperus enervis.

100
90
% germination

80

observed germination
fitted germination

70
60
50
40

r2 = 0.952762

30
20
10
0

1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 76 79 82
Days

Figure A6.3c
Figure 3c. Comparison of actual and fitted germination for the
'camphor' treatment of Cyperus enervis.

A6-3

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r = 0.971517

20
10
0
1

11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43
Days

Figure Figure
A6.4a 4a. Comparison of actual andfitted germination for the 'control'
treatment of Helichrysum bracteatum.

100
observed germination
fitted germination

90

% germination

80
70
60

r2 = 0.953791

50
40
30
20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43
Days

Figure A6.4b
Figure 4b. Comparison of actual and fitted germination for the 'cineole'
treatment of Helichrysum bracteatum.

100
observed germination
fitted germination

90
% germination

80
70
60
50

r2 = 0.973723

40
30
20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43
Days

Figure A6.4c
Figure 4c. Comparison of actual and fitted germination for the
'camphor' treatment of Helichrysum bracteatum.

A6-4

100
observed germination

90

fitted germination

% germination

80
70
60
50

r2 = 0.961131

40

r2 = 0.861131

30
20
10
0
1

9 13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85 89
Days

Figure A6.5a
Figure 5a. Comparison of actual and fitted germination for the 'control'
treatment of Lomandra longifolia.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30

rr22 == 0.862647
0.962647

20
10
0
1

9 13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85 89
Days

Figure Figure
A6.5b5b. Comparison of actual and fitted germination for the 'cineole'
treatment of Lomandra longifolia.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r2 = 0.993254
0.893254

10
0
1

9 13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85 89
Days

Figure A6.5c
Figure 5c. Comparison of actual andfitted germination for the
'camphor' treatment of Lomandra longifolia.

A6-5

100
observed germination

90

fitted germination

% germination

80
70
60
50

0.999343
r2 = 1.000057

40
30
20
10
0
1

11

13

15

17 19
Days

21

23

25

27

29

31

33

31

33

31

33

Figure Figure
A6.6a 6a. Comparison of actual and fitted germination for the 'control'
treatment of Oplismenus aemulus.

100
observed germination

90

fitted germination

% germination

80
70
60

r2 = 0.976489

50
40
30
20
10
0
1

11

13

15

17

19

21

23

25

27

29

Days

6b. Comparison of actual and fitted germination for the 'cineole'


FigureFigure
A6.6b

treatment of Oplismanus aemulus.

100

observed germination

90

fitted germination

% germination

80
70

r2 = 0.973359

60
50
40
30
20
10
0
1

11

13

15

17

19

21

23

Days

25

27

Figure A6.6c
Figure 6c. Comparison of actual and fitted germination for the
'camphor' treatment of Oplismenus aemulus.

A6-6

29

100
observed germination
fitted germination

90
% germination

80
70
60
50
40
2

r = 0.966393

30
20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days

Figure Figure
A6.7a7a. Comparison of actual and fitted germination for the 'control'
treatment of Ottochloa gracillima.

100
observed germination
fitted germination

90

% germination

80
70
60
50
40

r2 = 0.973744

30
20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days

FigureFigure
A6.7b7b. Comparison of actual and fitted germination for the 'cineole'
treatment of Ottochloa gracillima.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40

r2 = 0.980993

30
20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47
Days

Figure 7c. Comparison of actual and fitted germination for the


Figure A6.7c

'camphor' treatment of Ottochloa gracillima.

A6-7

100
observed germination

90

fitted germination

% germination

80
70
60
50
40

r = 0.9888381

30
20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days

Figure Figure
A6.8a8a. Comparison of actual and fitted germination for the 'control'
treatment of *Paspalum dilatatum.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30
20

r2 = 0.953132

10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days

Figure 8b. Comparison of actual and fitted germination for the


Figure A6.8b

'cineole' treatment of *Paspalum dilatatum.

100
observed germination
fitted germination

90
% germination

80
70
60

r2 = 1.062037

50
40
30

0.937363

20
10
0
1

9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45
Days

Figure A6.8c
Figure 8c. Comparison of actual andfitted germination for the
'camphor' treatment of *Paspalum dilatatum.

A6-8

100
90

r2 = 0.999643

80
% germination

70
60
50
40
30
20

observed germination

10

fitted germination

0
1

10 11 12 13 14 15 16 17 18 19 20 21
Days

Figure 9a. Comparison of actual and fitted germination for the 'control'
Figure A6.9a Comparison
of actual
and fitted germination
for the control
treatment
of *Pennesetum
clandestinum.
treatment of *Pennisetum clandestinum.

100
90
80

r2 = 0.959021

% germination

70
60
50
40
30
20

observed germination
fitted germination

10
0
1

10 11 12 13 14 15 16 17 18 19 20 21
Days

Figure

Figure 9b. Comparison of actual and fitted germination for the


A6.9b Comparison
of actualofand fitted germination for the cineole
'cineole' treatment
*Pennesetum clandestinum.
treatment of *Pennisetum clandestinum.

100
90

% germination

80
70
60
50

r2 = 0.964918

40
30
20

observed germination

10

fitted germination

0
1

10 11 12 13 14 15 16 17 18 19 20
Days

Figure

Figure 9c. Comparison of actual and fitted germination for the


A6.9c Comparison
of actual and
fitted germination
for the camphor
'camphor' treatment
of *Pennesetum
clandestinum.
treatment of *Pennisetum clandestinum.

A6-9

100
90

% germination

80

r = 0.987555

70
60
50
40
30
20

observed germination
fitted germination

10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25
Days

Figure A6.10a
Figure 10a. Comparison of actual and fitted germination for the
'control' treatment of Pratia purpurascens

100
90

% germination

80

r2 = 0.974188

70
60
50
40
30
20

observed germination
fitted germination

10
0
1

2 3

6 7

8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25
Days

Figure A6.10b
Figure 10b. Comparison of actual and fitted germination for the
'cineole' treatment of Pratia purpurascens.

100
90

% germination

80
70
60

r2 = 0.952142

50
40
30
20

observed germination
fitted germination

10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25
Days

Figure A6.10c
Figure 10c. Comparison of actual and fitted germination for the
'camphor' treatment of Pratia purpurascens.

A6-10

100
observed germination
fitted germination

90

% germination

80
70
60
50
40
30

r2 = 0.984786

20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
Days

Figure A6.11a
Figure 11a. Comparison of actual and fitted germination for the
'control' treatment of *Setaria sphaecelata.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40

r2 = 0.962518

30
20
10
0
1

10 11 12 13 14 15 16 17 18 19 20 21 22 23

Days

Figure 11b. Comparison of actual and fitted germination for the


Figure A6.11b

'cineole' treatment of *Setaria sphaecelata.

100
observed germination

90

fitted germination

% germination

80

r2 = 0.978009

70
60
50
40
30
20
10
0
1

10 11 12 13 14 15 16 17 18 19 20 21 22 23
Days

Figure A6.11c
Figure 11c. Comparison of actual and fitted germination for the
'camphor' treatment of *Setaria sphaecelata.

A6-11

100

observed germination

90

fitted germination

r2 = 0.998932

% germination

80
70
60
50
40
30
20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22
Days

Figure 12a. Comparison of actual and fitted germination for the


Figure A6.12a

'control' treatment of Viola hederacea.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40

r2 = 0.988413

30
20
10
0
1

9 10 11 12 13 14 15 16 17 18 19 20 21 22
Days

Figure 12b. Comparison of actual and fitted germination for the


Figure A6.12b

'cineole' treatment of Viola hederacea.

100
observed germination
fitted germination

90

% germination

80
70

r2 = 0.984958

60
50
40
30
20
10
0
1

10 11 12 13 14 15 16 17 18 19 20 21 22
Days

Figure A6.12c
Figure 12c. Comparison of actual and fitted germination for the
'camphor' treatment of Viola hederacea.

A6-12

Appendix 7.

Non-Linear Regressions - Vines


using the Richards function.

100
90

observed germination
fitted germination

% germination

80
70
60
50

r = 0.980419

40
30
20
10
0

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51 53 55 57
Days

Figure Figure
A7.1a1a. Comparison of actual and fitted germination for the 'control'
treatment of *Anredera cordifolia.
100
90

% germination

80

observed germination
fitted germination

70
60
50

r2 = 0.981533

40
30
20
10
0

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51 53 55 57
Days

Figure Figure
A7.1b1b. Comparison of actual and fitted germination for the 'cineole'
treatment of *Anredera cordifolia.
100
90

% germination

80

observed germination
fitted germination

70
60
50
40

r2 = 0.9656

30
20
10
0

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51 53 55 57
Days

Figure A7.1c
Figure 1c. Comparison of actual and fitted germination for the
'camphor' treatment of *Anredera cordifolia.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40

r2 = 0.974138

30
20
10
0
1

10

11

12

13

14

15

16

17

18

19

17

18 19

17

18

Days

FigureFigure
A7.2a2a. Comparison of actual and fitted germination for the 'contol'
treatment of *Cardiospermum grandiflorum.

100
observed germination

90

fitted germination

% germination

80
70
60
50
40
30

r2 = 0.98348

20
10
0
1

10

11

12

13

14 15

16

Days

FigureFigure
A7.2b2b. Comparison of actual and fitted germination for the 'cineole'
treatment of *Cardiospermum grandiflorum.

100

observed germination
fitted germination

90

% germination

80
70
60

r2 = 0.98764

50
40
30
20
10
0
1

10

11

12

13

14

Days

15

Figure A7.2c
Figure 2c. Comparison of actual and fitted germination for the
'camphor' treatment of *Cardiospermum grandiflorum.

A7-2

16

19

100
observed germination

90

fitted germination

% germination

80
70
60
50

r2 = 0.992703

40
30
20
10
0
1

13 17 21 25 29 33 37 41 45 49 53 57 61 65 69 73 77 81 85
Days

Figure 3a. Comparison of actual and fitted germination for the 'contol'
Figure A7.3a

treatment of Cissus hypoglauca.

100
90

observed germination
fitted germination

% germination

80
70
60

r2 = 0.998384

50
40
30
20
10
0

1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 76 79 82 85
Days

FigureFigure
A7.3b3b. Comparison of actual and fitted germination for the 'cineole'
treatment of Cissus hypoglauca.

90
80

observed germination
fitted germination

% germination

70
60
50
40

r2 = 0.995808

30
20
10
0

1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 76 79 82 85
Days

Figure A7.3c
Figure 3c. Comparison of actual and fitted germination for the
'camphor' treatment of Cissus hypoglauca.

Note: Geitonoplesium cymosum not assessed due to low germination of 2 percent in the control, 7 percent
in the cineole treatment and 0 percent in the camphor treatment.

A7-3

Appendix 8.

Modelled data from the Richards function


for the germination of all life-forms.

Species
Acacia melanoxylon

Algae culture 1

Algae culture 2

Algae culture 3

Allocasuarina littoralis

Allocasuarina torulosa

Alpinea caerulea

*Anredera cordifolia

Araucaria cunninghamii

Callistemon viminalis

*Cardiospermum
grandiflorum

*Cassia colluteoides

*Chloris gayana

*Cinnamomum camphora
(from soil)

Cissus hypoglauca

Treatment

Asymptote

MidInflection
Point

Days to MidInflection

Germination
Rate (GR)

Natural log
(ln) Days to
Mid-Inflection

control

50

18.26

10.28

0.440

2.33

cineole

59

25.80

18.77

0.590

2.93

camphor

73

26.90

13.34

0.430

2.59

control

100

36.77

7.51

0.700

2.02

cineole

101

37.26

38.48

0.270

3.65

camphor

116

42.54

73.56

0.150

4.30

control

100

58.50

13.23

-2.086

2.58

cineole

100

53.49

33.71

-1.660

3.52

camphor

120

44.29

68.65

0.119

4.23

control

100

85.90

15.80

-0.913

2.76

cineole

100

61.49

54.40

-0.360

4.00

camphor

102

47.96

71.46

1.130

4.27

control

43

15.99

4.48

1.330

1.50

cineole

42

15.59

4.22

2.230

1.44

camphor

43

15.96

4.19

2.100

1.43

control

19

8.02

12.77

1.140

2.55

cineole

12

4.26

13.89

0.240

2.63

camphor

2.84

11.23

3.790

2.42

control

52

18.96

52.50

0.098

3.96

cineole

10

3.71

50.45

0.235

3.92

camphor

45

16.69

48.67

0.170

3.89

control

76

28.14

21.78

0.220

3.08

cineole

64

26.67

33.36

0.200

3.51

camphor

72

26.38

32.67

0.100

3.49

control

32

11.72

7.79

0.900

2.05

cineole

33

12.13

7.38

0.866

2.00

camphor

32

13.47

8.18

1.700

2.10

control

71

26.12

5.00

1.560

1.61

cineole

49

18.14

7.00

1.110

1.95

camphor

72

26.44

5.00

1.300

1.61

control

60

42.09

16.50

-0.700

2.80

cineole

61

22.49

10.56

0.580

2.36

camphor

78

36.70

12.24

3.350

2.50

control

36

13.92

5.92

2.170

1.78

cineole

38

17.08

7.03

3.950

1.95

camphor

38

13.98

5.60

1.510

1.72

control

42

15.42

3.17

1.970

1.15

cineole

40

22.97

4.12

-4.090

1.42

camphor

35

12.79

3.74

0.950

1.32

control

28

13.21

21.51

2.730

3.07

cineole

16

5.88

23.28

0.270

3.15

camphor

6.70

92.05

-0.040

4.52

control

76

28.23

63.58

0.120

4.15

cineole

65

23.91

80.00

0.080

4.38

camphor

87

31.86

71.90

0.110

4.28

Commersonia bartramia

Cupaniopsis parvifolia

Cyperus enervis

Eucalyptus intermedia

Eucalyptus microcorys

Eucalyptus saligna

Ficus coronata

Ficus macrophylla

Geitonoplesium cymosum

control

71

26.27

16.49

0.310

2.80

cineole

82

41.71

33.68

-4.230

3.52

camphor

48

21.04

41.24

0.281

3.72

control

92

33.80

10.10

1.290

2.31

cineole

95

34.84

10.45

1.360

2.35

camphor

89

32.64

9.76

1.220

2.28

control

13

7.32

46.18

-0.450

3.83

cineole

13

4.73

33.68

0.110

3.52

camphor

25

9.22

41.10

0.110

3.72

control

88

39.11

7.42

3.658

2.00

cineole

89

32.65

8.65

0.769

2.16

camphor

94

34.76

7.28

1.951

1.99

control

28

12.52

10.21

2.130

2.32

cineole

33

12.19

12.93

0.580

2.56

camphor

26

10.18

10.56

1.070

2.36

control

78

28.71

6.84

1.050

1.92

cineole

71

26.06

6.92

1.090

1.93

camphor

91

33.48

8.00

0.640

2.08

control

69

25.26

16.25

0.460

2.79

cineole

97

35.86

18.53

0.662

2.92

camphor

86

34.41

27.83

0.820

3.33

control

6.43

16.00

-0.710

2.77

cineole

13

11.33

17.00

-0.880

2.83

camphor

6.26

17.00

-0.610

2.83

control

0.77

68.14

0.120

4.22

cineole

4.22

124.90

-5.840

4.83

#NULL!

#NULL!

#NULL!

#NULL!

#NULL!

control

77

27.95

7.00

1.100

1.95

cineole

77

28.33

8.00

0.990

2.08

camphor

87

31.92

7.00

1.070

1.95

control

14

5.25

6.00

0.540

1.79

cineole

10

3.76

12.50

0.170

2.53

camphor

14

5.08

15.00

0.268

2.71

control

73

32.90

79.94

0.470

4.38

cineole

33

17.36

82.68

-0.810

4.41

camphor

73

38.56

94.84

-0.660

4.55

control

89

32.60

9.62

1.000

2.26

cineole

89

32.60

9.62

1.000

2.26

camphor

95

35.00

9.99

1.500

2.30

control

98

39.25

38.37

0.370

3.65

cineole

47

28.91

48.53

-0.690

3.88

camphor

104

41.12

41.21

0.316

3.72

control

62

22.87

46.97

0.120

3.85

cineole

94

34.50

57.64

0.090

4.05

camphor

88

42.85

79.25

1.470

4.37

control

33

14.47

15.40

0.738

2.73

cineole

34

12.46

12.70

0.230

2.54

camphor

22

8.00

19.10

0.130

2.95

camphor
Guioa semiglauca

Helichrysum bracteatum

Hymenosporum flavum

Lepiderema pulchella

*Ligustrum lucidum

Lomandra longifolia

Lophostemon confertus

A8-2

Macaranga tanarius

Mallotus philippensis

Omalanthus populifolius

Oplismenus aemulus

Ottochloa gracillima

*Paspalum dilatatum

Pennisetum clandestinum

Pratia purpurescens

*Setaria shaecelata var.


narok

Solanum capsicoides

Syzygium luehmannii

Syzygium paniculatum

Toona australis

Tristaniopsis laurina

Viola hederacea

control

59

21.88

18.00

0.560

2.89

cineole

81

29.90

18.00

0.440

2.89

camphor

66

24.33

17.00

0.350

2.83

control

3.88

91.00

-0.100

4.51

cineole

19

6.88

39.00

0.180

3.66

camphor

2.48

52.00

0.150

3.95

control

83

41.11

15.98

24.180

2.77

cineole

79

32.27

15.89

1.220

2.77

camphor

83

30.35

15.99

0.760

2.77

control

82

41.60

39.05

-4.380

3.66

cineole

112

41.16

32.29

0.120

3.47

camphor

99

46.74

29.63

0.900

3.39

control

16

5.72

13.00

0.220

2.56

cineole

16

5.93

9.73

0.650

2.28

camphor

16

6.45

14.53

0.490

2.68

control

57

26.01

25.05

0.760

3.22

cineole

54

42.11

43.36

-0.160

3.77

camphor

7.94

43.17

-0.660

3.77

control

97

35.53

8.69

0.770

2.16

cineole

72

40.56

12.19

-1.960

2.50

camphor

77

36.43

10.18

3.090

2.32

control

92

38.78

11.45

0.616

2.44

cineole

82

30.00

6.10

1.040

1.81

camphor

83

30.43

5.79

2.250

1.76

control

12

4.44

8.00

0.890

2.08

cineole

21

7.61

7.31

1.140

1.99

camphor

14

5.41

8.63

0.700

2.16

control

88

36.02

14.49

1.470

2.67

cineole

73

34.64

22.31

3.060

3.11

camphor

65

24.08

17.51

0.460

2.86

control

93

34.68

11.59

1.310

2.45

cineole

88

32.20

12.39

1.050

2.52

camphor

91

33.57

12.58

0.805

2.53

control

88

32.29

13.00

0.530

2.56

cineole

87

32.10

12.00

0.700

2.48

camphor

78

28.64

13.00

0.580

2.56

control

88

45.00

13.00

-11.700

2.56

cineole

84

33.65

11.00

0.950

2.40

camphor

77

28.26

11.00

0.850

2.40

control

19

6.95

9.57

0.770

2.26

cineole

17

6.29

10.73

0.790

2.37

camphor

14

5.13

9.80

1.350

2.28

control

97

35.53

8.69

0.770

2.16

cineole

72

40.56

12.19

-1.960

2.50

camphor

77

36.43

10.18

3.090

2.32

Note: Lantana camara is not included in this analysis due to low germination of 2% in the control, 0%
cineole treatment and 6% camphor treatment.

A8-3

Appendix 9.

Results of the Deviance Tests (loglikelihood ratio)

applied to the modeled data for the germination of all life-forms.


Table A.9.1 Deviance test applied to the asymptote of germination i.e. the number of
seeds germinated.

Likelihood ratio test was 1196.390-1194.844 = 1.546 = chisquare on 2 df, p = 0.462.

Table A.9.2 Deviance test applied to the natural log (ln) of the Days to MidInflection i.e. time elapsed till germination began to slow.
lnDaysMidInfij ~ N(XB, )
lnDaysMidInfij = 0ijcons
0ij = 2.827(0.120) + u0j + e0ij
[u0j] ~ N(0, u) : u = [0.620(0.137)]
[e0ij] ~ N(0, e) : e = [0.113(0.117)]
-2*loglikelihood(IGLS Deviance) = 220.425(137 of 138 cases in use)

lnDaysMidInfij ~ N(XB, )
lnDaysMidInfij = 0ijcons + 0.169(0.066)Treat_2ij + 0.234(0.067)Treat_3ij
0ij = 2.694(0.126) + u0j + e0ij
[u0j] ~ N(0, u) : u = [0.628(0.138)]
[e0ij] ~ N(0, e) : e = [0.101(0.015)]
-2*loglikelihood(IGLS Deviance) = 207.920(137 of 138 cases in use)

Likelihood ratio test: 220.425-207.920 = 12.505 = chisquare on 2 df, p = 0.002.

Table A.9.3 Deviance test applied to the mid-inflection point of germination i.e. the
number of seeds germinated at the point where germination begins to slow.

Likelihood ratio test: 1011.261-1009.060 = 2.201 = chisquare on 2 df, p = 0.333.


Table A.4 Deviance test applied to the germination rate (GR).

Likelihood ratio test: 655.324-652.799 = 2.525 = chisquare on 2 df, p = 0.283.


A9-2

Appendix 11.

T-test comparisons of treatments

for shoot length, radicle length, leaf number and leaf area
of each species at completion of the germination triali

Acacia melanoxylon - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

40.60869565

35.83673469

Mean

40.60869565

40.28846154

Mean

35.83673469

40.28846154

Variance

128.2879227

90.38945578

Variance

128.2879227

180.3661388

Variance

90.38945578

180.3661388

46

49

46

52

49

52

Observations
Pooled Variance

108.7274236

Hypothesized Mean difference

2.22916639

1.66140353

Observations
Pooled Variance

df

0.014105474

t CritiCamphorl one-tail

155.954475

Hypothesized Mean difference

93

t Stat
P(T<=t) one-tail

Pooled Variance

df

Observations

136.7410803

Hypothesized Mean difference

96

df

99

t Stat

0.126688217

t Stat

P(T<=t) one-tail

0.449726071

P(T<=t) one-tail

-1.912132706
0.029375073

t CritiCamphorl one-tail

1.660882845

t CritiCamphorl one-tail

1.660391717

P(T<=t) two-tail

0.028210948

P(T<=t) two-tail

0.899452142

P(T<=t) two-tail

0.058750145

t CritiCamphorl two-tail

1.985799827

t CritiCamphorl two-tail

1.984985829

t CritiCamphorl two-tail

1.984217306

P(T<=t) two tail is

0.028210948

P(T<=t) two tail is

0.899452142

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Cineole

P(T<=t) two tail is

0.058750145

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in shoot length between Control and Camphor

*No Sig difference exists in shoot length between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-2

* Mean shoot length sig decreased in Cineole below Control

Radicle
Length
Acacia melanoxylon - Radical
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

32.39130435

30.51020408

Variance

51.93236715

115.880102

46

49

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat
P(T<=t) one-tail
t CritiCamphorl one-tail

84.93764967
0

32.78846154

Mean

Variance

51.93236715

161.1896682

Variance

46

52

Observations

Hypothesized Mean difference


df

0.99420737

t Stat

0.161350851
1.66140353

Cineole

32.39130435

Pooled Variance

93

Camphor

Mean

109.9753083

Observations
Pooled Variance

Hypothesized Mean difference

96

df

-0.187103704

t Stat

Camphor

30.51020408

32.78846154

115.880102

161.1896682

49

52

139.2213937
0
99
-0.969814866

P(T<=t) one-tail

0.425987093

P(T<=t) one-tail

0.167251108

t CritiCamphorl one-tail

1.660882845

t CritiCamphorl one-tail

1.660391717

P(T<=t) two-tail

0.322701702

P(T<=t) two-tail

0.851974186

P(T<=t) two-tail

0.334502216

t CritiCamphorl two-tail

1.985799827

t CritiCamphorl two-tail

1.984985829

t CritiCamphorl two-tail

1.984217306

P(T<=t) two tail is

0.322701702

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Cineole

P(T<=t) two tail is

0.851974186

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Camphor

P(T<=t) two tail is

0.334502216

value > 0.05 - null hypothesis (Ho) accepted

*No Sig difference exists in rad. length between Cineole and Camphor

Acacia melanoxylon - Phyllode Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean

Control

1.826086957

1.795918367

Mean

1.61352657

1.040816327

Variance

46

49

Variance
Observations
Pooled Variance

Cineole

1.317934186

Hypothesized Mean difference

Cineole

Camphor

1.403846154

Mean

1.795918367

1.403846154

1.61352657

0.951357466

Variance

1.040816327

0.951357466

46

52

49

52

1.261749234

Hypothesized Mean difference

93

Camphor

1.826086957

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

Pooled Variance

df

Observations

0.994731459

Hypothesized Mean difference

96

df

99

t Stat

0.128003927

t Stat

1.857125373

t Stat

1.974477381

P(T<=t) one-tail

0.449211128

P(T<=t) one-tail

0.033179619

P(T<=t) one-tail

0.025556657

t CritiCamphorl one-tail

1.660882845

t CritiCamphorl one-tail

1.660391717

t CritiCamphorl one-tail

1.66140353

P(T<=t) two-tail

0.898422255

P(T<=t) two-tail

0.066359238

P(T<=t) two-tail

0.051113313

t CritiCamphorl two-tail

1.985799827

t CritiCamphorl two-tail

1.984985829

t CritiCamphorl two-tail

1.984217306

P(T<=t) two tail is

0.898422255

P(T<=t) two tail is

0.066359238

value > 0.05 - null hypothesis (Ho) accepted

0.051113313

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf No. between Control and Camphor

* No Sig difference exists in leaf No. between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-3

* No Sig difference exists in leaf No. between Control and Cineole

P(T<=t) two tail is

Acacia melanoxylon - Phyllode Area

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean difference
df

Cineole

Control

66.7173913

56.18367347

Mean

3790.829469

2160.944728

Variance

46

49

2949.598635

Observations
Pooled Variance

Hypothesized Mean difference

93

df

Camphor

Cineole

Camphor

66.7173913

48.62745098

Mean

56.18367347

48.62745098

3790.829469

2524.278431

Variance

2160.944728

2524.278431

46

51

49

51

3124.22366

Observations
Pooled Variance

Hypothesized Mean difference

95

df

2346.319066
0
98

t Stat

0.944746675

t Stat

1.591637682

t Stat

P(T<=t) one-tail

0.173618134

P(T<=t) one-tail

0.057394181

P(T<=t) one-tail

0.218688089

t CritiCamphorl one-tail

1.66140353

0.77981993

t CritiCamphorl one-tail

1.661051101

t CritiCamphorl one-tail

1.660550879

P(T<=t) two-tail

0.347236269

P(T<=t) two-tail

0.114788362

P(T<=t) two-tail

0.437376178

t CritiCamphorl two-tail

1.985799827

t CritiCamphorl two-tail

1.985249583

t CritiCamphorl two-tail

1.984467417

P(T<=t) two tail is

0.347236269

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Cineole

P(T<=t) two tail is

0.114788362

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Camphor

P(T<=t) two tail is

0.437376178

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Cineole and Camphor

Allocasuarina littoralis - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

16.81818182

24.13043478

Mean

16.81818182

18.11627907

Mean

24.13043478

18.11627907

Variance

70.91774892

72.24927536

Variance

70.91774892

18.05758583

Variance

72.24927536

18.05758583

22

46

22

43

46

43

Observations
Pooled Variance

71.82560786

Hypothesized Mean difference

t Stat

Pooled Variance

df

Observations

35.67764019

Hypothesized Mean difference

66
-3.328490732

t Stat

Pooled Variance

df

Observations

46.08777007

Hypothesized Mean difference

63

df

-0.829083094

t Stat

87
4.176379552

P(T<=t) one-tail

0.000715579

P(T<=t) one-tail

0.205093838

P(T<=t) one-tail

3.50642E-05

t CritiCamphorl one-tail

1.668270215

t CritiCamphorl one-tail

1.669402536

t CritiCamphorl one-tail

1.662556315

P(T<=t) two-tail

0.001431157

P(T<=t) two-tail

0.410187677

P(T<=t) two-tail

7.01284E-05

t CritiCamphorl two-tail

1.996563697

t CritiCamphorl two-tail

1.998341759

t CritiCamphorl two-tail

1.987609721

P(T<=t) two tail is

0.001431157

P(T<=t) two tail is

0.410187677

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in shoot length between Control and Cineole

P(T<=t) two tail is

7.01284E-05

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in shoot length between Control and Camphor

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in shoot length between Cineole and Camphor


* Mean shoot length is increased in Cineole above Camphor

A11-4

* Mean shoot length is increased in Cineole above Control

Radicle
Length
Length
Allocasuarina littoralis - Radical
t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

6.590909091

7.326086957

Mean

6.590909091

7.348837209

Mean

7.326086957

7.348837209

Variance

11.20562771

9.735748792

Variance

11.20562771

12.85160576

Variance

9.735748792

12.85160576

22

46

22

43

46

43

Observations
Pooled Variance
Hypothesized Mean difference
df

10.20343754

Observations
Pooled Variance

Hypothesized Mean difference

66

df

12.30294641

Observations
Pooled Variance

Hypothesized Mean difference

63

df

11.2399556
0
87

t Stat

-0.88788142

t Stat

-0.82435133

t Stat

P(T<=t) one-tail

0.188914439

P(T<=t) one-tail

0.206425072

P(T<=t) one-tail

0.487276422

t CritiCamphorl one-tail

1.668270215

t CritiCamphorl one-tail

1.669402536

t CritiCamphorl one-tail

1.662556315

P(T<=t) two-tail

0.377828878

P(T<=t) two-tail

0.412850144

P(T<=t) two-tail

0.974552843

t CritiCamphorl two-tail

1.996563697

t CritiCamphorl two-tail

1.998341759

t CritiCamphorl two-tail

1.987609721

P(T<=t) two tail is

0.377828878

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in rad length between Control and Cineole

P(T<=t) two tail is

0.412850144

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in rad length between Control and Camphor

-0.031990577

P(T<=t) two tail is

0.974552843

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in rad length between Cineole and Camphor

Allocasuarina littoralis - No. of Phyllode Branches

0 values - stats not possible

No dif in phyllode no.between Control and Cineole, Control and Camphor, Cineole and Camphor

0 values - stats not possible

0 values - stats not possible

0 values - stats not possible

No dif in phyllode length between Control and Cineole

No dif in phyllode length between Control and Camphor

No dif in phyllode length between Cineole and Camphor

A11-5

Allocasuarina littoralis - Phyllode Length

Allocasuarina torulosa - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

51.30769231

38.6

Variance

126.0641026

239.6

13

10

Observations
Pooled Variance

174.7223443

Hypothesized Mean difference

Cineole

51.30769231

52.66666667

Mean

Variance

126.0641026

363.3333333

Variance

13

12

Observations

239.5406912

Hypothesized Mean difference

21

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

df

52.66666667

239.6

363.3333333

10

12

Observations
Pooled Variance

307.6533333

Hypothesized Mean difference

23

df

t Stat

2.285599986

t Stat

0.016387371

P(T<=t) one-tail

0.414160811

P(T<=t) one-tail

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

1.713870006

t CritiCamphorl one-tail

1.724718004

P(T<=t) two-tail

0.032774743

P(T<=t) two-tail

0.828321623

P(T<=t) two-tail

0.075758761

t CritiCamphorl two-tail

2.079614205

t CritiCamphorl two-tail

2.068654794

t CritiCamphorl two-tail

2.085962478

0.032774743

P(T<=t) two tail is

0.828321623

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in shoot length between Control and Cineole

t Stat

20

P(T<=t) one-tail

P(T<=t) two tail is

-0.219338015

Camphor
38.6

-1.873008147
0.03787938

P(T<=t) two tail is

0.075758761

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* no sig dif exists in shoot length between Control and Camphor

* no sig dif exists in shoot length between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-6

* Mean shoot length reduced in Cineole compared to Control

Length
Allocasuarina torulosa - Radical
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

40.38461538

32.5

Variance

72.75641026

156.9444444

13

10

Observations
Pooled Variance
Hypothesized Mean difference
df

108.8369963
0

44.16666667

Mean

Variance

72.75641026

199.2424242

Variance

13

12

Observations

Hypothesized Mean difference


df

t Stat

1.796801069

t Stat

P(T<=t) one-tail

0.043383263

P(T<=t) one-tail

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

P(T<=t) two-tail

0.086766525

t CritiCamphorl two-tail

2.079614205

P(T<=t) two tail is

0.086766525

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in rad length between Control and Cineole

Cineole

40.38461538

Pooled Variance

21

Camphor

Mean

133.2497213

Observations
Pooled Variance

Hypothesized Mean difference

23

df

-0.818439447

t Stat

0.2107527

Camphor
32.5

44.16666667

156.9444444

199.2424242

10

12

180.2083333
0
20
-2.029731713

P(T<=t) one-tail

0.027950037

1.713870006

t CritiCamphorl one-tail

1.724718004

P(T<=t) two-tail

0.421505401

P(T<=t) two-tail

0.055900074

t CritiCamphorl two-tail

2.068654794

t CritiCamphorl two-tail

2.085962478

P(T<=t) two tail is

0.421505401

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in rad length between Control and Camphor

P(T<=t) two tail is

0.055900074

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in rad length between Cineole and Camphor

Allocasuarina torulosa - No. of Phyllode Branches

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

2.076923077

0.7

Variance

4.076923077

2.455555556

13

10

Observations
Pooled Variance

3.382051282

Hypothesized Mean difference

Cineole

2.076923077

2.416666667

Mean

Variance

4.076923077

3.356060606

Variance

13

12

Observations

3.732162765

Hypothesized Mean difference

21

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

df

2.416666667

2.455555556

3.356060606

10

12

Observations
Pooled Variance

2.950833333

Hypothesized Mean difference

23

df

1.780030085

t Stat

P(T<=t) one-tail

0.044773418

P(T<=t) one-tail

0.332271189

P(T<=t) one-tail

0.015071605

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

1.713870006

t CritiCamphorl one-tail

1.724718004

P(T<=t) two-tail

0.089546837

P(T<=t) two-tail

0.664542378

P(T<=t) two-tail

t CritiCamphorl two-tail

2.079614205

t CritiCamphorl two-tail

2.068654794

t CritiCamphorl two-tail

0.089546837

P(T<=t) two tail is

0.664542378

value > 0.05 - null hypothesis (Ho) accepted

* no sig dif exists in phyllode branching between Control and Cineole

t Stat

20

t Stat

P(T<=t) two tail is

-0.439302099

Camphor
0.7

-2.333957355

0.03014321
2.085962478

P(T<=t) two tail is

0.03014321

value > 0.05 - null hypothesis (Ho) accepted

* no sig dif exists in phyllode branching between Control and Camphor

value < 0.05 - null hypothesis (Ho) rejected

*Sig dif exists in phylode branching between Control and Camphor

A11-7

t-Test: Two-Sample Assuming Equal Variances

* Phyllode branching is reduced in Cineole below Camphor

Allocasuarina torulosa - Phyllode Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Control

1.9

26.33333333

11.65555556

13

10

Observations
Pooled Variance

Cineole

20.04285714

Hypothesized Mean difference

Mean
Variance

1.646226682

t Stat

P(T<=t) one-tail

0.057301065

P(T<=t) one-tail

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

P(T<=t) two-tail
t CritiCamphorl two-tail

0.11460213

14.90909091

13

12

20.86956522

t CritiCamphorl two-tail

0.11460213

* No sig dif exists between Control and Cineole

11.65555556

14.90909091

10

12

Observations

13.445
0

df

t Stat

1.713870006
1
2.068654794

value > 0.05 - null hypothesis (Ho) accepted

Variance

20
-1.974515618

P(T<=t) one-tail

0.031146492

t CritiCamphorl one-tail

1.724718004

P(T<=t) two-tail

0.062292984

t CritiCamphorl two-tail

2.085962478

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists betweenControl and Camphor

Camphor
1.9

Hypothesized Mean difference

0.5

P(T<=t) two tail is

Mean

Pooled Variance

23

P(T<=t) two-tail

2.079614205

P(T<=t) two tail is

26.33333333

df

t Stat

Cineole
5

Hypothesized Mean difference

21

Camphor
5

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

0.062292984

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists betweenCineole and Camphor

Alpinea caerulea - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

22.03921569

16.22222222

Mean

22.03921569

23.29032258

Mean

16.22222222

23.29032258

Variance

33.75843137

129.4102564

Variance

33.75843137

15.97990481

Variance

129.4102564

15.97990481

51

27

51

62

27

62

Observations
Pooled Variance

66.48142415

Hypothesized Mean difference

t Stat

t Stat

0.00183779
1.665150648
0.00367558
1.991675163

P(T<=t) two tail is

0.00367558

49.87863058

Hypothesized Mean difference

111

df

-1.351253538

t Stat

87
-4.340384163

P(T<=t) one-tail

0.089680956

P(T<=t) one-tail

1.91203E-05

t CritiCamphorl one-tail

1.658697784

t CritiCamphorl one-tail

1.662556315

P(T<=t) two-tail

0.179361912

P(T<=t) two-tail

3.82406E-05

t CritiCamphorl two-tail

1.981566129

t CritiCamphorl two-tail

1.987609721

P(T<=t) two tail is

0.179361912

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Cineole

Observations
Pooled Variance

df

2.997561399

P(T<=t) two-tail
t CritiCamphorl two-tail

23.98825011

Hypothesized Mean difference

76

P(T<=t) one-tail
t CritiCamphorl one-tail

Pooled Variance

df

Observations

P(T<=t) two tail is

3.82406E-05

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in shoot length between Control and Camphor

value < 0.05 - null hypothesis (Ho) rejected

*Sig difference exists in shoot length between Cineole and Camphor


* Mean shoot length sig increased in Camphor above Cineole

A11-8

* Mean shoot length sig decreased in Cineole below Control

Radicle
Length
Length
Alpinea caerulea - Radical
t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

41.31372549

21.03571429

Mean

41.31372549

40.32258065

Mean

21.03571429

40.32258065

Variance

279.6196078

174.9246032

Variance

279.6196078

91.69751454

Variance

174.9246032

91.69751454

51

28

51

62

28

62

Observations
Pooled Variance

242.9083724

Hypothesized Mean difference


df
t Stat
P(T<=t) one-tail
t CritiCamphorl one-tail
P(T<=t) two-tail
t CritiCamphorl two-tail

Pooled Variance

Hypothesized Mean difference

77

df

5.531649761
2.0973E-07
1.664884621
4.1946E-07
1.991256795

P(T<=t) two tail is

4.1946E-07

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Cineole


* Mean rad length sig decreased in Cineole below Control

Observations

176.3471061

Observations
Pooled Variance

Hypothesized Mean difference

111

df

117.2330986
0
88

t Stat

0.394815932

t Stat

P(T<=t) one-tail

0.346868181

P(T<=t) one-tail

5.31051E-12

t CritiCamphorl one-tail

1.658697784

t CritiCamphorl one-tail

1.662353952

P(T<=t) two-tail

0.693736363

P(T<=t) two-tail

t CritiCamphorl two-tail

1.981566129

t CritiCamphorl two-tail

P(T<=t) two tail is

0.693736363

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Camphor

-7.823303072

1.0621E-11
1.987291398

P(T<=t) two tail is

1.0621E-11

value < 0.05 - null hypothesis (Ho) rejected

*Sig difference exists in rad. length between Cineole and Camphor


* Mean rad length sig increased in Camphor above Cineole

Alpinea caerulea - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Control

1.94

0.892857143

Mean

0.465714286

1.876984127

Variance

50

28

Observations
Pooled Variance

Cineole

0.967086466

Hypothesized Mean difference

Camphor

Mean

0.892857143

1.758064516

0.465714286

0.219196192

Variance

1.876984127

0.219196192

50

62

28

62

0.329008798

Observations
Pooled Variance

df

4.51117836

Cineole

1.758064516

Hypothesized Mean difference

76

t Stat

Camphor
1.94

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

0.727835672

Hypothesized Mean difference

110

df

88

t Stat

1.668727758

t Stat

P(T<=t) one-tail

1.15437E-05

P(T<=t) one-tail

0.049007653

P(T<=t) one-tail

1.23287E-05

t CritiCamphorl one-tail

1.665150648

t CritiCamphorl one-tail

1.658822839

t CritiCamphorl one-tail

1.662353952

P(T<=t) two-tail

2.30873E-05

P(T<=t) two-tail

0.098015306

P(T<=t) two-tail

2.46575E-05

t CritiCamphorl two-tail

1.991675163

t CritiCamphorl two-tail

1.981766218

t CritiCamphorl two-tail

1.987291398

P(T<=t) two tail is

2.30873E-05

P(T<=t) two tail is

0.098015306

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf No. between Control and Cineole

-4.454071136

P(T<=t) two tail is

2.46575E-05

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf No. between Control and Camphor

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf No. between Cineole and Camphor


* Mean leaf no. sig reduced in Cineole below Camphor

A11-9

* Mean leaf no. sig reduced in Cineole below Control

Alpinea caerulea - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

131.9215686

70.64285714

Mean

131.9215686

155.0967742

Mean

70.64285714

155.0967742

Variance

2255.073725

13138.97884

Variance

2255.073725

1562.646219

Variance

13138.97884

1562.646219

51

28

51

62

28

62

Observations
Pooled Variance
Hypothesized Mean difference
df

6071.507985

Observations
Pooled Variance

1874.550501

Hypothesized Mean difference

77

df

Observations
Pooled Variance

Hypothesized Mean difference

111

df

3.343581376

t Stat

P(T<=t) one-tail

0.000639686

P(T<=t) one-tail

0.002751124

P(T<=t) one-tail

6.78207E-07

t CritiCamphorl one-tail

1.664884621

t CritiCamphorl one-tail

1.658697784

t CritiCamphorl one-tail

1.662353952

P(T<=t) two-tail

0.001279371

P(T<=t) two-tail

0.005502249

P(T<=t) two-tail

1.35641E-06

t CritiCamphorl two-tail

1.991256795

t CritiCamphorl two-tail

1.981566129

t CritiCamphorl two-tail

1.987291398

0.001279371

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

t Stat

0
88

t Stat

P(T<=t) two tail is

-2.831501093

5114.475545

0.005502249

value < 0.05 - null hypothesis (Ho) rejected

-5.1864817

P(T<=t) two tail is

1.35641E-06

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Control and Cineole

* Sig difference exists in leaf area between Control and Camphor

* Sig difference exists in leaf area between Cineole and Camphor

* Mean leaf area sig reduced in Cineole compared to Control

* Mean leaf area sig increased in Camphor above Control

* Mean leaf area sig. increased in Camphor above Cineole

*Anredera cordifolia - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

22.22535211

17.94915254

Mean

22.22535211

20.1

Variance

70.43420523

67.32495617

Variance

70.43420523

48.09130435

71

59

71

70

Observations
Pooled Variance

69.02532675

Hypothesized Mean difference

Pooled Variance

df

Observations

59.34312494

Hypothesized Mean difference

128

17.94915254

20.1

Variance

67.32495617

48.09130435

59

70

Observations
Pooled Variance

df

56.87517683

Hypothesized Mean difference

139

Camphor

Mean

df

127

t Stat

2.921710854

t Stat

1.638001605

t Stat

P(T<=t) one-tail

0.002058183

P(T<=t) one-tail

0.051841564

P(T<=t) one-tail

0.054535004

t CritiCamphorl one-tail

1.656844688

t CritiCamphorl one-tail

1.655889719

t CritiCamphorl one-tail

1.656940185

P(T<=t) two-tail

0.004116366

P(T<=t) two-tail

0.103683128

P(T<=t) two-tail

0.109070009

t CritiCamphorl two-tail

1.978669388

t CritiCamphorl two-tail

1.977177817

t CritiCamphorl two-tail

1.978819455

P(T<=t) two tail is

0.004116366

P(T<=t) two tail is

0.103683128

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Cineole

-1.613720816

P(T<=t) two tail is

0.109070009

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in shoot length between Control and Camphor

*No Sig difference exists in shoot length between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-10

* Mean shoot length sig decreased in Cineole below Control

Radicle
Length
*Anredera cordifolia - Radical
Length
t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

Cineole

Control

28.5915493

31.13333333

Mean

317.2736419

338.5242938

Variance

71

60

326.9929323

Observations
Pooled Variance

Hypothesized Mean difference

129

df

-0.801564762

t Stat

Camphor

Cineole

Camphor

28.5915493

35.10144928

Mean

31.13333333

35.10144928

317.2736419

423.2101449

Variance

338.5242938

423.2101449

71

69

60

69

369.4742376

Observations
Pooled Variance

Hypothesized Mean difference

138

df

-2.003419687

t Stat

383.8678991
0
127
-1.147357531

P(T<=t) one-tail

0.212139398

P(T<=t) one-tail

0.023545207

P(T<=t) one-tail

0.126695365

t CritiCamphorl one-tail

1.656751465

t CritiCamphorl one-tail

1.655971573

t CritiCamphorl one-tail

1.656940185

P(T<=t) two-tail

0.424278795

P(T<=t) two-tail

0.047090415

P(T<=t) two-tail

0.253390731

t CritiCamphorl two-tail

1.978523869

t CritiCamphorl two-tail

1.977305146

t CritiCamphorl two-tail

1.978819455

P(T<=t) two tail is

0.424278795

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Cineole

P(T<=t) two tail is

0.047090415

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Camphor


* Mean rad length sig increased in Camphor compared to Control

P(T<=t) two tail is

0.253390731

value > 0.05 - null hypothesis (Ho) accepted

*No Sig difference exists in rad. length between Cineole and Camphor

*Anredera cordifolia - Leaf Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

1.591549296

1.55

Variance

0.245070423

0.251694915

71

60

Observations
Pooled Variance

0.248100229

Hypothesized Mean difference

1.442857143

Mean

Variance

0.245070423

0.250310559

Variance

71

70

Observations

0.247671641

t Stat

0.475684758

t Stat

P(T<=t) one-tail

0.317551762

P(T<=t) one-tail

t CritiCamphorl one-tail

1.656751465

t CritiCamphorl one-tail

P(T<=t) two-tail

0.635103524

P(T<=t) two-tail

t CritiCamphorl two-tail

1.978523869

t CritiCamphorl two-tail

0.635103524

0.0391382
1.655889719
0.0782764
1.977177817

0.0782764

value > 0.05 - null hypothesis (Ho) accepted

70

0.250948661
0
128

t Stat

1.215692539

P(T<=t) one-tail

0.113169727

t CritiCamphorl one-tail

1.656844688

P(T<=t) two-tail

0.226339455

t CritiCamphorl two-tail

1.978669388

P(T<=t) two tail is

0.226339455

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf No. between Control and Camphor

* No Sig difference exists in leaf No. between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-11

* No Sig difference exists in leaf No. between Control and Cineole

0.250310559

60

df

1.773856757

P(T<=t) two tail is

1.442857143

0.251694915

Hypothesized Mean difference

139

Camphor
1.55

Observations
Pooled Variance

df

P(T<=t) two tail is

Cineole

1.591549296

Hypothesized Mean difference

129

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

*Anredera cordifolia - Leaf Area


t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

150.7183099

138.4166667

Mean

150.7183099

136.6666667

Mean

138.4166667

136.6666667

Variance

5544.805231

4802.179379

Variance

5544.805231

5878.578431

Variance

4802.179379

5878.578431

71

60

71

69

60

69

Observations
Pooled Variance
Hypothesized Mean difference
df

5205.154648

Observations
Pooled Variance

Hypothesized Mean difference

129

df

5709.273185

Observations
Pooled Variance

Hypothesized Mean difference

138

df

5378.519029
0
127

t Stat

0.972333971

t Stat

1.100085406

t Stat

0.135179865

P(T<=t) one-tail

0.166352057

P(T<=t) one-tail

0.136605105

P(T<=t) one-tail

0.446341841

t CritiCamphorl one-tail

1.656751465

t CritiCamphorl one-tail

1.655971573

t CritiCamphorl one-tail

1.656940185

P(T<=t) two-tail

0.332704115

P(T<=t) two-tail

P(T<=t) two-tail

0.892683683

t CritiCamphorl two-tail

1.978523869

t CritiCamphorl two-tail

t CritiCamphorl two-tail

1.978819455

P(T<=t) two tail is

0.332704115

value > 0.05 - null hypothesis (Ho) accepted

*No Sig difference exists in leaf area between Control and Cineole

P(T<=t) two tail is

0.27321021
1.977305146

0.27321021

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Camphor

P(T<=t) two tail is

0.892683683

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Cineole and Camphor

Araucaria cunninghamii - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Cineole

Pooled Variance

44.03333333

Mean

57.30555556

36.99885057

Variance

28

30

46.78958333

Hypothesized Mean difference

-0.435810475

t Stat

0.332324811

P(T<=t) one-tail

t CritiCamphorl one-tail

1.672522103

t CritiCamphorl one-tail

P(T<=t) two-tail

0.664649622

t CritiCamphorl two-tail

2.003239388

P(T<=t) two tail is

0.664649622

44.03333333

42.75862069

57.30555556

108.1896552

Variance

36.99885057

108.1896552

28

29

30

29

83.21018809

Observations
Pooled Variance

71.96977213

Hypothesized Mean difference

55

df

0.203314897

t Stat

0.41981963

57
0.576993406

P(T<=t) one-tail

0.283107935

1.673033694

t CritiCamphorl one-tail

1.672028702

P(T<=t) two-tail

0.839639259

P(T<=t) two-tail

t CritiCamphorl two-tail

2.004044291

t CritiCamphorl two-tail

P(T<=t) two tail is

0.839639259

value > 0.05 - null hypothesis (Ho) accepted

0.56621587
2.002466317

P(T<=t) two tail is

0.56621587

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in shoot length between Control and Camphor

* No sig difference exists in shoot length between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-12

* No sig difference exists in shoot length between Control and Cineole

Camphor

Mean

df

P(T<=t) one-tail

Cineole

42.75862069

Hypothesized Mean difference

56

Camphor
43.25

Observations
Pooled Variance

df
t Stat

Control

43.25

Observations

t-Test: Two-Sample Assuming Equal Variances

Radicle
Length
Length
Araucaria cunninghamii - Radical
t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat
P(T<=t) one-tail

Cineole

Control

38.75

41.83333333

Mean

112.2685185

64.62643678

Variance

28

30

87.59672619

Observations
Pooled Variance

Hypothesized Mean difference

56

df

-1.253724713
0.107575426

Camphor

Cineole

Camphor

38.75

33.79310345

Mean

41.83333333

33.79310345

112.2685185

258.3128079

Variance

64.62643678

258.3128079

28

29

30

29

186.6183386

Observations
Pooled Variance

159.7706191

Hypothesized Mean difference

55

df

57

t Stat

1.369535465

t Stat

2.442607171

P(T<=t) one-tail

0.088199037

P(T<=t) one-tail

0.008852305

t CritiCamphorl one-tail

1.672522103

t CritiCamphorl one-tail

1.673033694

t CritiCamphorl one-tail

1.672028702

P(T<=t) two-tail

0.215150852

P(T<=t) two-tail

0.176398074

P(T<=t) two-tail

0.017704611

t CritiCamphorl two-tail

2.003239388

t CritiCamphorl two-tail

2.004044291

t CritiCamphorl two-tail

2.002466317

P(T<=t) two tail is

0.215150852

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in rad. length between Control and Cineole

P(T<=t) two tail is

0.176398074

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in rad. length between Control and Camphor

0.017704611

value < 0.05 - null hypothesis (Ho) rejected

*Sig difference exists in rad. length between Cineole and Camphor


* Mean rad length greater in Cineole than Camphor

Araucaria cunninghamii - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

15.60714286

11.3

Variance

191.8029101

91.18275862

28

30

Observations
Pooled Variance

139.6960459

Hypothesized Mean difference

Cineole

15.60714286

15.4137931

Variance

191.8029101

135.7512315

28

29

Observations

163.2675101

Hypothesized Mean difference

56

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

Variance

15.4137931

91.18275862

135.7512315

30

29

113.0760436

Hypothesized Mean difference

55

Camphor
11.3

Observations
Pooled Variance

df

Mean

df

57

t Stat

1.386828844

t Stat

0.057112915

t Stat

P(T<=t) one-tail

0.085494914

P(T<=t) one-tail

0.477331118

P(T<=t) one-tail

0.071452587

t CritiCamphorl one-tail

1.672522103

t CritiCamphorl one-tail

1.673033694

t CritiCamphorl one-tail

1.672028702

P(T<=t) two-tail

0.170989829

P(T<=t) two-tail

0.954662237

P(T<=t) two-tail

0.142905175

t CritiCamphorl two-tail

2.003239388

t CritiCamphorl two-tail

2.004044291

t CritiCamphorl two-tail

2.002466317

P(T<=t) two tail is

0.170989829

P(T<=t) two tail is

0.954662237

value > 0.05 - null hypothesis (Ho) accepted

P(T<=t) two tail is

0.142905175

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in leaf No. between Control and Camphor

* No sig difference exists in leaf No. between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-13

* No sig difference exists in leaf No. between Control and Cineole

-1.485561939

Araucaria cunninghamii - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

4.037037037

2.933333333

Mean

4.037037037

4.931034483

Mean

2.933333333

4.931034483

Variance

12.11396011

6.547126437

Variance

12.11396011

11.78078818

Variance

6.547126437

11.78078818

27

30

27

29

30

29

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

9.178720539

Observations
Pooled Variance

Hypothesized Mean difference

55

df

1.37330344

t Stat

11.94120429

Observations
Pooled Variance

9.118047994

Hypothesized Mean difference

54

df

-0.967384403

t Stat

57
-2.540464924

P(T<=t) one-tail

0.087615374

P(T<=t) one-tail

0.168832372

P(T<=t) one-tail

0.006909592

t CritiCamphorl one-tail

1.673033694

t CritiCamphorl one-tail

1.673565748

t CritiCamphorl one-tail

1.672028702

P(T<=t) two-tail

0.175230748

P(T<=t) two-tail

0.337664744

P(T<=t) two-tail

0.013819183

t CritiCamphorl two-tail

2.004044291

t CritiCamphorl two-tail

2.004881026

t CritiCamphorl two-tail

2.002466317

P(T<=t) two tail is

0.175230748

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in leaf area between Control and Cineole

P(T<=t) two tail is

0.337664744

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in leaf area between Control and Camphor

0.013819183

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Cineole and Camphor


* Mean leaf area greater in Camphor than Cineole

Callistemon viminalis - Shoot Length

Cineole treatment predated (analysis voided)

t-Test: Two-Sample Assuming Equal Variances


Cineole treatment predated (analysis voided)
Control

Camphor

Mean

12.90769231

14.59649123

Variance

18.77259615

14.60213033

65

57

Observations
Pooled Variance

16.82637877

Hypothesized Mean difference

df

120

t Stat

-2.26880308

P(T<=t) one-tail

0.012533573

t CritiCamphorl one-tail

1.657649591

P(T<=t) two-tail

0.025067145

t CritiCamphorl two-tail

1.979929038

P(T<=t) two tail is

0.025067145

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in shoot length between Control and Camphor

A11-14

* Mean shoot length Camphor increased above Control

Radicle
Length
Length
Callistemon viminalis - Radical

Cineole treatment predated (analysis voided)

t-Test: Two-Sample Assuming Equal Variances

Cineole treatment predated (analysis voided)

Control

Camphor

Mean

13.06153846

12.66666667

Variance

52.96490385

64.44047619

65

57

Observations
Pooled Variance

58.32017094

Hypothesized Mean difference

df

120

t Stat

0.284944915

P(T<=t) one-tail

0.388088766

t CritiCamphorl one-tail

1.657649591

P(T<=t) two-tail

0.776177531

t CritiCamphorl two-tail

1.979929038

P(T<=t) two tail is

0.776177531

value > 0.05 - null hypothesis (Ho) accepted

*Sig dif in Rad length between Control and Camphor


* Mean rad length Camphor is reduced below Control

Callistemon viminalis - Leaf Number


Cineole treatment predated (analysis voided)

t-Test: Two-Sample Assuming Equal Variances

Cineole treatment predated (analysis voided)

Control
Mean
Variance

1.719298246

0.860863095

0.884085213

64

57

Observations
Pooled Variance

0.871791151

Hypothesized Mean difference

df
t Stat

Camphor

4.109375

119
14.05530744

P(T<=t) one-tail

2.4047E-27

t CritiCamphorl one-tail

1.65775873

P(T<=t) two-tail

4.80939E-27

t CritiCamphorl two-tail

1.980097295

P(T<=t) two tail is

4.80939E-27

value < 0.05 - null hypothesis (Ho) rejected

* sig dif in leaf no. exists between Control and Camphor

A11-15

* Mean leaf No reduced in Camphor

Callistemon viminalis - Leaf Area

Cineole treatment predated (analysis voided)

t-Test: Two-Sample Assuming Equal Variances

Cineole treatment predated (analysis voided)

Control

Camphor

Mean

3.661290323

1.456140351

Variance

7.604706504

1.502506266

62

57

Observations
Pooled Variance

4.683995279

Hypothesized Mean difference

df
t Stat

117
5.552514465

P(T<=t) one-tail

8.93446E-08

t CritiCamphorl one-tail

1.657981556

P(T<=t) two-tail

1.78689E-07

t CritiCamphorl two-tail

1.98044745

P(T<=t) two tail is

1.78689E-07

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif in leaf area between Control and Camphor


* Mean leaf area is reduced in Camphor

*Cardiospermum grandiflorum - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

75.22580645

77.29411765

Mean

75.22580645

92.27941176

Mean

77.29411765

92.27941176

Variance

416.9317821

464.6117647

Variance

416.9317821

579.4282265

Variance

464.6117647

579.4282265

62

51

62

68

51

68

Pooled Variance

438.4092518

Hypothesized Mean difference

t Stat

Pooled Variance

df

Observations

501.9885147

Hypothesized Mean difference

111
-0.522537831

t Stat

Pooled Variance

df

Observations

530.3613625

Hypothesized Mean difference

128

df

-4.334589175

t Stat

117
-3.512735067

P(T<=t) one-tail

0.301168641

P(T<=t) one-tail

1.46417E-05

P(T<=t) one-tail

0.000315634

t CritiCamphorl one-tail

1.658697784

t CritiCamphorl one-tail

1.656844688

t CritiCamphorl one-tail

1.657981556

P(T<=t) two-tail

0.602337283

P(T<=t) two-tail

2.92834E-05

P(T<=t) two-tail

0.000631268

t CritiCamphorl two-tail

1.981566129

t CritiCamphorl two-tail

1.978669388

t CritiCamphorl two-tail

P(T<=t) two tail is

0.602337283

P(T<=t) two tail is

2.92834E-05

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in shoot length between Control and Cineole

1.98044745

P(T<=t) two tail is

0.000631268

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Camphor

*Sig difference exists in shoot length between Cineole and Camphor

* Mean shoot length sig increased in Camphor above Control

* Mean shoot length sig increased in Camphor above Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-16

Observations

*Cardiospermum grandiflorum - Radical


Length
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

35.40322581

47.80487805

Mean

35.40322581

51.39705882

Mean

47.80487805

51.39705882

Variance

357.6216288

700.0609756

Variance

357.6216288

323.7653644

Variance

700.0609756

323.7653644

62

41

62

68

41

68

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

493.2411721

Observations
Pooled Variance

Hypothesized Mean difference

101

df

-2.774080434

t Stat

339.8999904

Observations
Pooled Variance

Hypothesized Mean difference

128

df

-4.940325982

t Stat

464.4366209
0
107
-0.843001004

P(T<=t) one-tail

0.003297586

P(T<=t) one-tail

1.19484E-06

P(T<=t) one-tail

0.200554196

t CritiCamphorl one-tail

1.660080216

t CritiCamphorl one-tail

1.656844688

t CritiCamphorl one-tail

1.659218469

P(T<=t) two-tail

0.006595173

P(T<=t) two-tail

2.38968E-06

P(T<=t) two-tail

0.401108391

t CritiCamphorl two-tail

1.983730726

t CritiCamphorl two-tail

1.978669388

t CritiCamphorl two-tail

1.982384674

P(T<=t) two tail is

0.006595173

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

2.38968E-06

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Cineole

* Sig difference exists in rad. length between Control and Camphor

* Mean rad length sig increased in Cineole compared to Control

* Mean rad length sig increased in Camphor compared to Control

P(T<=t) two tail is

0.401108391

value > 0.05 - null hypothesis (Ho) accepted

*No Sig difference exists in rad. length between Cineole and Camphor

*Cardiospermum grandiflorum - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

2.225806452

2.254901961

Variance

0.374405077

0.55372549

62

51

Observations
Pooled Variance

0.455180038

Hypothesized Mean difference

t Stat

2.367647059

Mean

Variance

0.374405077

0.415057068

Variance

62

68

Observations

0.395683853

t Stat

P(T<=t) one-tail

0.409983738

P(T<=t) one-tail

t CritiCamphorl one-tail

1.658697784

t CritiCamphorl one-tail

P(T<=t) two-tail

0.819967476

P(T<=t) two-tail

t CritiCamphorl two-tail

1.981566129

t CritiCamphorl two-tail

P(T<=t) two tail is

0.819967476

51

68

0.474317077
0

t Stat

0.10071
1.656844688
0.20142
1.978669388

0.20142

-0.883749847

P(T<=t) one-tail

0.189321817

t CritiCamphorl one-tail

1.657981556

P(T<=t) two-tail

0.378643635

t CritiCamphorl two-tail

P(T<=t) two tail is

117

1.98044745

P(T<=t) two tail is

0.378643635

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf No. between Control and Camphor

* No Sig difference exists in leaf No. between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-17

* No Sig difference exists in leaf No. between Control and Cineole

0.415057068

df

-1.284118322

value > 0.05 - null hypothesis (Ho) accepted

2.367647059

0.55372549

Hypothesized Mean difference

128

Camphor

2.254901961

Observations
Pooled Variance

df

-0.228126674

Cineole

2.225806452

Hypothesized Mean difference

111

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

*Cardiospermum grandiflorum - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Mean

321.1451613

250.3921569

Mean

321.1451613

194.761194

Variance

108625.7655

42235.64314

Variance

108625.7655

24788.21483

62

51

62

67

Observations
Pooled Variance
Hypothesized Mean difference
df

78720.30496

Observations
Pooled Variance

Hypothesized Mean difference

111

df

65056.64467

250.3921569

194.761194

Variance

42235.64314

24788.21483

51

67

Observations
Pooled Variance

Hypothesized Mean difference

127

Camphor

Mean

df

32308.65807
0
116

t Stat

1.333961248

t Stat

2.811800764

t Stat

P(T<=t) one-tail

0.092473543

P(T<=t) one-tail

0.002854551

P(T<=t) one-tail

0.049258082

t CritiCamphorl one-tail

1.658697784

t CritiCamphorl one-tail

1.656940185

t CritiCamphorl one-tail

1.658095243

P(T<=t) two-tail

0.184947086

P(T<=t) two-tail

0.005709101

P(T<=t) two-tail

0.098516164

t CritiCamphorl two-tail

1.981566129

t CritiCamphorl two-tail

1.978819455

t CritiCamphorl two-tail

1.980624802

P(T<=t) two tail is

0.184947086

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Cineole

P(T<=t) two tail is

0.005709101

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Control and Camphor


* Mean leaf area sig reduced in Camphor compared to Control

P(T<=t) two tail is

1.665476857

0.098516164

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Cineole and Camphor

*Cassia coleutioides - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

57.24137931

55.80645161

Variance

274.2610837

506.827957

29

31

Observations
Pooled Variance

394.554294

Hypothesized Mean difference

Cineole

57.24137931

70.74074074

Mean

Variance

274.2610837

253.2763533

Variance

29

27

Observations

264.1573246

Hypothesized Mean difference

58

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

df

70.74074074

506.827957

253.2763533

31

27

Observations
Pooled Variance

389.1075696

Hypothesized Mean difference

54

df

0.279628126

t Stat

P(T<=t) one-tail

0.390378814

P(T<=t) one-tail

0.001510963

P(T<=t) one-tail

0.002842956

t CritiCamphorl one-tail

1.671553491

t CritiCamphorl one-tail

1.673565748

t CritiCamphorl one-tail

1.672522103

P(T<=t) two-tail

0.780757628

P(T<=t) two-tail

0.003021927

P(T<=t) two-tail

0.005685912

t CritiCamphorl two-tail

2.001715984

t CritiCamphorl two-tail

2.004881026

t CritiCamphorl two-tail

2.003239388

0.780757628

P(T<=t) two tail is

0.003021927

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in shoot length between Control and Cineole

t Stat

56

t Stat

P(T<=t) two tail is

-3.105766235

Camphor

55.80645161

-2.876063348

P(T<=t) two tail is

0.005685912

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in shoot length between Control and Camphor

* Sig dif exists in shootlength between Cineole and Camphor

* Mean shoot length increased in Camphor compared to Control

* Mean shoot length of Cineole reduced below Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-18

t-Test: Two-Sample Assuming Equal Variances

Radicle
Length
*Cassia coleutioides - Radical
Length

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

43.48275862

38.38709677

Variance

167.1157635

442.311828

29

31

Observations
Pooled Variance
Hypothesized Mean difference
df

309.4585555
0

Cineole

43.48275862

40.03703704

Mean

Variance

167.1157635

361.1908832

Variance

29

27

Observations
Pooled Variance
Hypothesized Mean difference

58

Camphor

Mean

df

260.5593397

Observations
Pooled Variance

Hypothesized Mean difference

54

df

Camphor

38.38709677

40.03703704

442.311828

361.1908832

31

27

404.6485322
0
56

t Stat

1.121252886

t Stat

0.798203685

t Stat

P(T<=t) one-tail

0.133400591

P(T<=t) one-tail

0.214124378

P(T<=t) one-tail

0.378256117

t CritiCamphorl one-tail

1.671553491

t CritiCamphorl one-tail

1.673565748

t CritiCamphorl one-tail

1.672522103

P(T<=t) two-tail

0.266801183

P(T<=t) two-tail

0.428248755

P(T<=t) two-tail

0.756512235

t CritiCamphorl two-tail

2.001715984

t CritiCamphorl two-tail

2.004881026

t CritiCamphorl two-tail

2.003239388

P(T<=t) two tail is

0.266801183

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in rad length between Control and Cineole

P(T<=t) two tail is

0.428248755

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in rad length between Control and Camphor

-0.311586023

P(T<=t) two tail is

0.756512235

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in rad length between Cineole and Camphor

*Cassia coleutioides - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

1.551724138

1.258064516

Mean

1.551724138

1.444444444

Mean

1.258064516

1.444444444

Variance

0.256157635

0.264516129

Variance

0.256157635

0.256410256

Variance

0.264516129

0.256410256

29

31

29

27

31

27

Observations
Pooled Variance

0.260480994

Hypothesized Mean difference

Pooled Variance

df

Observations

0.256279268

Hypothesized Mean difference

58

Pooled Variance

df

Observations

0.260752688

Hypothesized Mean difference

54

df

56

t Stat

2.227206244

t Stat

0.792405816

t Stat

P(T<=t) one-tail

0.014914486

P(T<=t) one-tail

0.215795813

P(T<=t) one-tail

0.085538224

t CritiCamphorl one-tail

1.671553491

t CritiCamphorl one-tail

1.673565748

t CritiCamphorl one-tail

1.672522103

P(T<=t) two-tail

0.029828972

P(T<=t) two-tail

0.431591625

P(T<=t) two-tail

0.171076448

t CritiCamphorl two-tail

2.001715984

t CritiCamphorl two-tail

2.004881026

t CritiCamphorl two-tail

2.003239388

P(T<=t) two tail is

0.029828972

P(T<=t) two tail is

0.431591625

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in leaf no. between Control and Cineole

-1.386543424

P(T<=t) two tail is

0.171076448

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in leaf no between Control and Camphor

* No sig dif exists in leaf no between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-19

* Mean leaf no reduced in Cineole compared to Control

*Cassia coleutioides - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Cineole
63.58064516

Mean

2474.455172

2171.051613

Variance

30

31

Observations
Pooled Variance

Control

94.6

2320.182176

Hypothesized Mean difference

Camphor

Mean

63.58064516

62.55555556

2474.455172

1264.717949

Variance

2171.051613

1264.717949

30

27

31

27

1902.579394

Hypothesized Mean difference

59

Cineole

62.55555556

Observations
Pooled Variance

df

Camphor
94.6

Pooled Variance

df

Observations

Hypothesized Mean difference

55

df

1750.25384
0
56

t Stat

2.514480515

t Stat

2.769404757

t Stat

0.093080786

P(T<=t) one-tail

0.007333653

P(T<=t) one-tail

0.003820756

P(T<=t) one-tail

0.463085798

t CritiCamphorl one-tail

1.671091923

t CritiCamphorl one-tail

1.673033694

t CritiCamphorl one-tail

1.672522103

P(T<=t) two-tail

0.014667307

P(T<=t) two-tail

0.007641512

P(T<=t) two-tail

0.926171597

t CritiCamphorl two-tail

2.000997483

t CritiCamphorl two-tail

2.004044291

t CritiCamphorl two-tail

2.003239388

P(T<=t) two tail is

0.014667307

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.007641512

value < 0.05 - null hypothesis (Ho) rejected

* sig dif exists in leaf area between Control and Cineole

* sig dif exists in leaf area between Control and Camphor

* Mean leaf area is reduced below Control

* Mean leaf area is reduced below Control

P(T<=t) two tail is

0.926171597

value > 0.05 - null hypothesis (Ho) accepted

* no sig dif exists in leaf area between Cineole and Camphor

*Cinnamomum camphora (fresh from tree) - Shoot Length

0 value for Control - stats not possible

0 value for Control - stats not possible

t-Test: Two-Sample Assuming Equal Variances

Cineole

*dif exists in shoot length between Control and Cineole

* dif exists in shoot length between Control and Camphor

* Mean shoot length Cineole increased above Control

* Mean shoot length Camphor increased above Control

Camphor

Mean

143.3870968

97.7173913

Variance

1210.957223

746.3807931

93

92

Observations
Pooled Variance

979.9383426

Hypothesized Mean difference

df
t Stat

183
9.921523357

P(T<=t) one-tail

3.88053E-19

t CritiCamphorl one-tail

1.653222625

P(T<=t) two-tail

7.76105E-19

t CritiCamphorl two-tail

1.973012331

P(T<=t) two tail is

0.452714098

value > 0.05 - null hypothesis (Ho) accepted

A11-20

* No sig dif exists in shoot length between Cineole and Camphor

Radicle
Length
Length
*Cinnamomum camphora (fresh from tree) - Radical
0 value for Control - stats not possible

0 value for Control - stats not possible

t-Test: Two-Sample Assuming Equal Variances

Cineole

Camphor

Mean

66.79569892

52.55434783

477.2730248

168.4036073

93

92

*dif exists between Control and Cineole

* dif exists between Control and Camphor

Variance

* Cineole increased above Control

* Camphor increased above Control

Observations
Pooled Variance

323.6822215

Hypothesized Mean difference

df
t Stat

183
5.383209654

P(T<=t) one-tail

1.11159E-07

t CritiCamphorl one-tail

1.653222625

P(T<=t) two-tail

2.22317E-07

t CritiCamphorl two-tail

1.973012331

P(T<=t) two tail is

2.22317E-07

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in rad length between Cineole and Camphor


* Mean Camphor rad length reduced below Cineole

*Cinnamomum camphora (fresh from tree) - Leaf Number

0 value for Control - stats not possible

0 value for Control - stats not possible

t-Test: Two-Sample Assuming Equal Variances

Cineole

Camphor

Mean

3.935483871

3.77173913

1.082748948

0.661610129

93

92

*dif in leaf no exists between Control and Cineole

* dif in leaf no exists between Control and Camphor

Variance

* Cineole increased above Control

* Camphor increased above Control

Observations
Pooled Variance

0.873330191

Hypothesized Mean difference

df

183

t Stat

1.191592185

P(T<=t) one-tail

0.117482134

t CritiCamphorl one-tail

1.653222625

P(T<=t) two-tail

0.234964268

t CritiCamphorl two-tail

1.973012331

P(T<=t) two tail is

0.234964268

A11-21

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in leaf No between Cineole and Camphor

*Cinnamomum camphora (fresh from tree) - Leaf Area

0 value for Control - stats not possible

0 value for Control - stats not possible

t-Test: Two-Sample Assuming Equal Variances

Cineole

Camphor

Mean

478.9784946

372.7826087

71987.69518

44700.28189

93

92

*dif in leaf area exists between Control and Cineole

* dif in leaf area exists between Control and Camphor

Variance

* Cineole increased above Control

* Camphor increased above Control

Observations
Pooled Variance
Hypothesized Mean Difference
df

58418.54431
0
183

t Stat

2.988008652

P(T<=t) one-tail

0.001596694

t Critical one-tail

1.653222625

P(T<=t) two-tail

0.003193388

t Critical two-tail

1.973012331

P(T<=t) two-tail

0.003193388

value < 0.05 - null hypothesis (Ho) rejected


* Sig dif exists in leaf area between Cineole and Camphor
*Cineole mean is larger than Camphor

*Cinnamomum camphora (from soil) - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

184.6153846

209

Mean

184.6153846

172.5

Variance

3481.089744

610

Variance

3481.089744

2491.666667

13

10

13

Observations
Pooled Variance

2250.622711

Hypothesized Mean difference

df
t Stat

Observations
Pooled Variance

3283.205128

Hypothesized Mean difference

21
-1.222003134

209

172.5

Variance

610

2491.666667

10

Observations
Pooled Variance

df

1080.416667

Hypothesized Mean difference

15

Camphor

Mean

df

12

t Stat

0.369798724

t Stat

1.876995385

P(T<=t) one-tail

0.117623232

P(T<=t) one-tail

0.358350691

P(T<=t) one-tail

0.042518512

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

1.753051038

t CritiCamphorl one-tail

1.782286745

P(T<=t) two-tail

0.235246464

P(T<=t) two-tail

0.716701383

P(T<=t) two-tail

0.085037025

t CritiCamphorl two-tail

2.079614205

t CritiCamphorl two-tail

2.131450856

t CritiCamphorl two-tail

2.178812792

P(T<=t) two tail is

0.235246464

P(T<=t) two tail is

0.716701383

value > 0.05 - null hypothesis (Ho) accepted

0.085037025

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in shoot length between Control and Camphor

* No sig dif exists in shoot length between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-22

* No sig dif exists in shoot length between Control and Cineole

P(T<=t) two tail is

Length
Radicle
Length
*Cinnamomum camphora (from soil) - Radical
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

105.3846154

56

Variance

3414.423077

371.1111111

13

10

Observations
Pooled Variance

2110.14652

Hypothesized Mean difference

93.75

Variance

3414.423077

256.25

13

Observations

Hypothesized Mean difference

21

Cineole

105.3846154

Pooled Variance

df

Camphor

Mean

df

2782.788462
0
15

Mean
Variance
Observations
Pooled Variance
Hypothesized Mean difference
df

Camphor
56

93.75

371.1111111

256.25

10

342.3958333
0
12

t Stat

2.555895703

t Stat

0.385735087

t Stat

P(T<=t) one-tail

0.009202986

P(T<=t) one-tail

0.352554215

P(T<=t) one-tail

0.002409556

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

1.753051038

t CritiCamphorl one-tail

1.782286745

P(T<=t) two-tail

0.018405971

P(T<=t) two-tail

P(T<=t) two-tail

0.004819113

t CritiCamphorl two-tail

2.079614205

t CritiCamphorl two-tail

t CritiCamphorl two-tail

2.178812792

P(T<=t) two tail is

0.018405971

value < 0.05 - null hypothesis (Ho) rejected

0.70510843
2.131450856

0.70510843

value > 0.05 - null hypothesis (Ho) accepted

P(T<=t) two tail is

-3.448408431

0.004819113

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif in rad length exists between Control and Cineole


* Mean rad length reduced in Cineole below Control

* No sig dif in rad length exists between Control and Camphor

* Sig dif exists in rad length between Cineole and Camphor


* Mean rad length reduced in Cineole compared to Camphor

*Cinnamomum camphora (from soil) - Leaf Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

10.92307692

12.9

Variance

21.24358974

25.87777778

13

10

Observations
Pooled Variance

23.22967033

Hypothesized Mean difference

t Stat

11.5

Variance

21.24358974

12.33333333

13

Observations

19.46153846

t Stat

Mean
Variance

11.5

25.87777778

12.33333333

10

22.49166667

Hypothesized Mean difference

15

df

-0.228721262

Camphor
12.9

Observations
Pooled Variance

df

-0.975160286

Cineole

10.92307692

Hypothesized Mean difference

21

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

12

t Stat

0.498980064

P(T<=t) one-tail

0.170290473

P(T<=t) one-tail

0.411087288

P(T<=t) one-tail

0.313407362

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

1.753051038

t CritiCamphorl one-tail

1.782286745

P(T<=t) two-tail

0.340580945

P(T<=t) two-tail

0.822174575

P(T<=t) two-tail

0.626814723

t CritiCamphorl two-tail

2.079614205

t CritiCamphorl two-tail

2.131450856

t CritiCamphorl two-tail

2.178812792

P(T<=t) two tail is


P(T<=t) two tail is

0.340580945

P(T<=t) two tail is

0.626814723

0.822174575
value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in leaf no between Control and Cineole

* No sig difference exists in leaf no between Cineole and Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-23

* No sig difference exists in leaf no between Control and Cineole

value > 0.05 - null hypothesis (Ho) accepted

*Cinnamomum camphora (from soil) - Leaf Area


t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

642.3846154

530.2

Variance

61904.75641

40760.4

13

10

Observations
Pooled Variance
Hypothesized Mean difference
df

52842.88938
0

Cineole

642.3846154

639.75

Variance

61904.75641

105426.9167

13

Observations
Pooled Variance
Hypothesized Mean difference

21

Camphor

Mean

df

70609.18846

Mean
Variance
Observations
Pooled Variance

Hypothesized Mean difference

15

df

Camphor

530.2

639.75

40760.4

105426.9167

10

56927.02917
0
12

t Stat

1.160239865

t Stat

0.017340588

t Stat

P(T<=t) one-tail

0.129486103

P(T<=t) one-tail

0.493196729

P(T<=t) one-tail

0.226357049

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

1.753051038

t CritiCamphorl one-tail

1.782286745

P(T<=t) two-tail

0.258972205

P(T<=t) two-tail

0.986393458

P(T<=t) two-tail

0.452714098

t CritiCamphorl two-tail

2.079614205

t CritiCamphorl two-tail

2.131450856

t CritiCamphorl two-tail

2.178812792

P(T<=t) two tail is

0.258972205

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference in leaf area exists between Control and Cineole

P(T<=t) two tail is

0.986393458

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference in leaf area exists between Control and Camphor

P(T<=t) two tail is

-0.776102531

0.452714098

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference in leaf area exists between Cineole and Camphor

Cissus hypoglauca - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

44.67605634

39.24615385

Mean

44.67605634

35.3375

Variance

155.5649899

197.7509615

Variance

155.5649899

128.277057

71

65

71

80

Observations
Pooled Variance

175.7135137

Hypothesized Mean difference

Pooled Variance

df

141.0968912

Hypothesized Mean difference

134

t Stat

Observations

2.38619454

35.3375

Variance

197.7509615

128.277057

65

80

Observations

159.370273

Hypothesized Mean difference

149

t Stat

39.24615385

Pooled Variance

df

df

143

t Stat

1.854134449

P(T<=t) one-tail

0.009210951

P(T<=t) one-tail

1.73883E-06

P(T<=t) one-tail

0.032889947

t CritiCamphorl one-tail

1.656303539

t CritiCamphorl one-tail

1.655143933

t CritiCamphorl one-tail

P(T<=t) two-tail

0.018421902

P(T<=t) two-tail

3.47765E-06

P(T<=t) two-tail

0.065779895

t CritiCamphorl two-tail

1.977823558

t CritiCamphorl two-tail

1.976013664

t CritiCamphorl two-tail

1.976691237

P(T<=t) two tail is

0.018421902

4.82177249

Camphor

Mean

P(T<=t) two tail is

3.47765E-06

value < 0.05 - null hypothesis (Ho) rejected

1.65558049

P(T<=t) two tail is

0.065779895

value < 0.05 - null hypothesis (Ho) rejected

value > 0.05 - null hypothesis (Ho) accepted

* Sig difference exists in shoot length between Control and Camphor

* Mean shoot length sig decreased in Cineole below Control

* Mean shoot length sig decreased in Camphor below Control

*No Sig difference exists in shoot length between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-24

* Sig difference exists in shoot length between Control and Cineole

Cissus hypoglauca - Radical


Length
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Mean

36.33802817

31.69230769

Mean

36.33802817

29.525

Variance

344.2555332

224.1538462

Variance

344.2555332

153.6196203

71

65

71

80

Observations
Pooled Variance
Hypothesized Mean difference
df

286.8935334

Observations
Pooled Variance

Hypothesized Mean difference

134

df

243.1801163

31.69230769

29.525

Variance

224.1538462

153.6196203

65

80

Observations
Pooled Variance

Hypothesized Mean difference

149

Camphor

Mean

df

185.1873857
0
143

t Stat

1.597750987

t Stat

2.679549892

t Stat

P(T<=t) one-tail

0.056226936

P(T<=t) one-tail

0.004100704

P(T<=t) one-tail

t CritiCamphorl one-tail

1.656303539

t CritiCamphorl one-tail

1.655143933

t CritiCamphorl one-tail

P(T<=t) two-tail

0.112453873

P(T<=t) two-tail

0.008201408

P(T<=t) two-tail

0.341821845

t CritiCamphorl two-tail

1.977823558

t CritiCamphorl two-tail

1.976013664

t CritiCamphorl two-tail

1.976691237

P(T<=t) two tail is

0.112453873

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Cineole

P(T<=t) two tail is

0.008201408

value < 0.05 - null hypothesis (Ho) rejected

0.95374559
0.170910923
1.65558049

P(T<=t) two tail is

0.341821845

value > 0.05 - null hypothesis (Ho) accepted

* Sig difference exists in rad. length between Control and Camphor

*Sig difference exists in rad. length between Cineole and Camphor

* Mean rad length sig decreased in Camphor compared to Control

* Mean rad length sig decreased in Camphor below Cineole

Cissus hypoglauca - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

0.985714286

0.738461538

Mean

0.985714286

0.75

Variance

0.217184265

0.321153846

Variance

0.217184265

0.240506329

70

65

70

80

Observations
Pooled Variance

0.26721474

Hypothesized Mean difference

Pooled Variance

df

Observations

0.229633205

Hypothesized Mean difference

133

0.738461538

0.75

Variance

0.321153846

0.240506329

65

80

Observations
Pooled Variance

df

0.276600323

Hypothesized Mean difference

148

Camphor

Mean

df

143

t Stat

2.776832824

t Stat

3.005501471

t Stat

P(T<=t) one-tail

0.003141019

P(T<=t) one-tail

0.001557651

P(T<=t) one-tail

t CritiCamphorl one-tail

1.656389941

t CritiCamphorl one-tail

1.655214419

t CritiCamphorl one-tail

P(T<=t) two-tail

0.006282039

P(T<=t) two-tail

0.003115301

P(T<=t) two-tail

0.895656984

t CritiCamphorl two-tail

1.977959982

t CritiCamphorl two-tail

1.976122803

t CritiCamphorl two-tail

1.976691237

P(T<=t) two tail is

0.006282039

P(T<=t) two tail is

0.003115301

value < 0.05 - null hypothesis (Ho) rejected

-0.131383019
0.447828492
1.65558049

P(T<=t) two tail is

0.895656984

value < 0.05 - null hypothesis (Ho) rejected

value > 0.05 - null hypothesis (Ho) accepted

* Sig difference exists in leaf No. between Control and Camphor


* Mean leaf no. sig reduced in Camphor below Control

* No Sig difference exists in leaf No. between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-25

* Sig difference exists in leaf No. between Control and Cineole


* Mean leaf no. sig reduced in Cineole below Control

Cissus hypoglauca - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean difference
df

Cineole

Control

224.8

104.1818182

Mean

32196.42319

17289.35105

Variance

70

66

24965.38073

Observations
Pooled Variance

Hypothesized Mean difference

134

df

Camphor

Cineole

224.8

111.3875

32196.42319

19603.15174

70

80

25474.3391

104.1818182

111.3875

Variance

17289.35105

19603.15174

66

80

Observations
Pooled Variance

Hypothesized Mean difference

148

df

0
144

t Stat

4.449336112

t Stat

8.97157E-06

P(T<=t) one-tail

1.30433E-05

P(T<=t) one-tail

0.375441108

t CritiCamphorl one-tail

1.656303539

t CritiCamphorl one-tail

1.655214419

t CritiCamphorl one-tail

1.655503183

P(T<=t) two-tail

1.79431E-05

P(T<=t) two-tail

2.60866E-05

P(T<=t) two-tail

0.750882217

t CritiCamphorl two-tail

1.977823558

t CritiCamphorl two-tail

1.976122803

t CritiCamphorl two-tail

1.976577551

1.79431E-05

value < 0.05 - null hypothesis (Ho) rejected

t Stat

18558.72782

P(T<=t) one-tail

P(T<=t) two tail is

4.34168034

Camphor

Mean

P(T<=t) two tail is

2.60866E-05

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Control and Cineole

* Sig difference exists in leaf area between Control and Camphor

* Mean leaf area sig reduced in Cineole compared to Control

* Mean leaf area sig reduced in Camphor compared to Control

P(T<=t) two tail is

-0.318083772

0.750882217

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Cineole and Camphor

Commersonia bartramia - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Control

26.5

24.03571429

Mean

16.92028986

28.51678141

Variance

70

84

Observations
Pooled Variance

Cineole

23.25258459

Hypothesized Mean difference

Cineole

Camphor

22.24390244

Mean

24.03571429

22.24390244

16.92028986

13.73902439

Variance

28.51678141

13.73902439

70

41

84

41

15.75285299

Hypothesized Mean difference

152

Camphor
26.5

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

Pooled Variance

df

Observations

23.71100677

Hypothesized Mean difference

109

df

123

t Stat

3.157793639

t Stat

5.452697734

t Stat

P(T<=t) one-tail

0.000958864

P(T<=t) one-tail

1.56126E-07

P(T<=t) one-tail

1.931496703
0.027860981

t CritiCamphorl one-tail

1.654939297

t CritiCamphorl one-tail

1.658954716

t CritiCamphorl one-tail

1.657335815

P(T<=t) two-tail

0.001917729

P(T<=t) two-tail

3.12252E-07

P(T<=t) two-tail

0.055721963

t CritiCamphorl two-tail

1.975695341

t CritiCamphorl two-tail

1.981966307

t CritiCamphorl two-tail

1.979437911

P(T<=t) two tail is

0.001917729

P(T<=t) two tail is

3.12252E-07

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.055721963

value < 0.05 - null hypothesis (Ho) rejectted

* Sig difference exists in shoot length between Control and Camphor


* Mean shoot length sig decreased in Camphor below Control

*No Sig difference exists in shoot length between Cineole and Camphor

A11-26

* Sig difference exists in shoot length between Control and Cineole


* Mean shoot length sig decreased in Cineole below Control

value > 0.05 - null hypothesis (Ho) accepted

Commersonia bartramia - Radical


Length
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

29.85714286

Mean

118.5507246

140.9913941

Variance

70

84

130.8045113

Hypothesized Mean difference

Observations
Pooled Variance

df
t Stat

Control

24

Observations
Pooled Variance

Cineole

t-Test: Two-Sample Assuming Equal Variances

Hypothesized Mean difference

152

df

-3.164482935

Camphor

Cineole

24

24.46341463

118.5507246

141.304878

70

41

126.9008727

29.85714286

24.46341463

Variance

140.9913941

141.304878

84

41

Observations
Pooled Variance

141.0933401

Hypothesized Mean difference

109

Camphor

Mean

df

123

t Stat

-0.20917843

t Stat

P(T<=t) one-tail

0.000938438

P(T<=t) one-tail

0.417349692

P(T<=t) one-tail

2.383484693
0.009339798

t CritiCamphorl one-tail

1.654939297

t CritiCamphorl one-tail

1.658954716

t CritiCamphorl one-tail

1.657335815

P(T<=t) two-tail

0.001876875

P(T<=t) two-tail

0.834699385

P(T<=t) two-tail

0.018679597

t CritiCamphorl two-tail

1.975695341

t CritiCamphorl two-tail

1.981966307

t CritiCamphorl two-tail

1.979437911

P(T<=t) two tail is

0.001876875

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Cineole


* Mean rad length increased in Cineole above Control

P(T<=t) two tail is

0.834699385

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Camphor

0.018679597

value < 0.05 - null hypothesis (Ho) rejected

*Sig difference exists in rad. length between Cineole and Camphor


* Mean rad length greater in Cineole than Camphor

Commersonia bartramia - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

1.171428571

Mean

1.171428571

0.87804878

Variance

0.173084886

Variance

0.173084886

0.109756098

70

84

70

41

Observations
Pooled Variance

0.078571429

Hypothesized Mean difference

df

Observations
Pooled Variance

0.149844964

Hypothesized Mean difference

152

0.87804878

Variance

0.109756098

84

41

Observations
Pooled Variance

df

0.03569304

Hypothesized Mean difference

109

Camphor

Mean

df

123

t Stat

3.779019944

t Stat

3.853794728

t Stat

3.388214869

P(T<=t) one-tail

0.000112798

P(T<=t) one-tail

9.84561E-05

P(T<=t) one-tail

0.000472782

t CritiCamphorl one-tail

1.654939297

t CritiCamphorl one-tail

1.658954716

t CritiCamphorl one-tail

1.657335815

P(T<=t) two-tail

0.000225597

P(T<=t) two-tail

0.000196912

P(T<=t) two-tail

0.000945564

t CritiCamphorl two-tail

1.975695341

t CritiCamphorl two-tail

1.981966307

t CritiCamphorl two-tail

1.979437911

P(T<=t) two tail is

0.000225597

P(T<=t) two tail is

0.000196912

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.000945564

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf No. between Control and Camphor

* Sig difference exists in leaf No. between Cineole and Camphor

* Mean leaf no. sig reduced in Camphor below Control

* Mean leaf no. reduced in Camphor below Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-27

* Sig difference exists in leaf No. between Control and Cineole


* Mean leaf no. reduced in Cineole compared to Control

Commersonia bartramia - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

4.385714286

1.535714286

Variance

5.660662526

2.90232358

70

84

Observations
Pooled Variance

4.154464286

Hypothesized Mean difference

1.536585366

Mean

Variance

5.660662526

1.954878049

Variance

70

41

Observations

4.300741617

Hypothesized Mean difference

152

Cineole

4.385714286

Pooled Variance

df

Camphor

Mean

df

Observations
Pooled Variance
Hypothesized Mean difference

109

df

Camphor

1.535714286

1.536585366

2.90232358

1.954878049

84

41

2.594211212
0
123

t Stat

8.640037903

t Stat

6.985860102

t Stat

P(T<=t) one-tail

3.61336E-15

P(T<=t) one-tail

1.17563E-10

P(T<=t) one-tail

0.498869791

t CritiCamphorl one-tail

1.654939297

t CritiCamphorl one-tail

1.658954716

t CritiCamphorl one-tail

1.657335815

P(T<=t) two-tail

7.22671E-15

P(T<=t) two-tail

2.35125E-10

P(T<=t) two-tail

0.997739582

t CritiCamphorl two-tail

1.975695341

t CritiCamphorl two-tail

1.981966307

t CritiCamphorl two-tail

1.979437911

P(T<=t) two tail is

7.22671E-15

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

2.35125E-10

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Control and Cineole

* Sig difference exists in leaf area between Control and Camphor

* Mean leaf area sig reduced in Cineole below Control

* Mean leaf area reduced in Camphor below Control

P(T<=t) two tail is

-0.002838782

0.997739582

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Cineole and Camphor

Cupaniopsis parvifolia - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

65.38383838

58.49484536

Variance

184.4430014

160.044244

99

97

Observations
Pooled Variance

172.3693895

Hypothesized Mean difference

Cineole

65.38383838

69.40625

Variance

184.4430014

293.8648026

99

96

Observations

238.3034735

Hypothesized Mean difference

194

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

Variance

69.40625

160.044244

293.8648026

97

96

226.6042077

Hypothesized Mean difference

193

df

3.672832324

t Stat

P(T<=t) one-tail

0.000155083

P(T<=t) one-tail

0.035222748

P(T<=t) one-tail

5.50858E-07

t CritiCamphorl one-tail

1.652747414

t CritiCamphorl one-tail

1.652788342

t CritiCamphorl one-tail

1.652870196

P(T<=t) two-tail

0.000310166

P(T<=t) two-tail

0.070445497

P(T<=t) two-tail

1.10172E-06

t CritiCamphorl two-tail

1.972266546

t CritiCamphorl two-tail

t CritiCamphorl two-tail

1.972462087

0.000310166

1.97233021

P(T<=t) two tail is

0.070445497

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif in shoot length exists between Control and Cineole

t Stat

191

t Stat

P(T<=t) two tail is

-1.819102757

Camphor

58.49484536

Observations
Pooled Variance

df

Mean

-5.034884424

P(T<=t) two tail is

1.10172E-06

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif in shoot length exists between Control and Camphor

value < 0.05 - null hypothesis (Ho) rejected

*Sig dif in shoot length exists between Cineole and Camphor


* Mean shoot length is sig reduced in Cineole compared to Camphor

A11-28

* Mean shoot length reduced in Cineole below Control

Cupaniopsis parvifolia - Radical


Radicle
Length
Length
t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

54.54545455

47.2371134

Variance

197.4953618

79.93277491

99

97

Observations
Pooled Variance
Hypothesized Mean difference
df

t-Test: Two-Sample Assuming Equal Variances

139.3200611
0

Cineole

54.54545455

45.3125

Variance

197.4953618

81.48026316

99

96

Observations
Pooled Variance
Hypothesized Mean difference

194

Camphor

Mean

df

140.3894842

Mean
Variance
Observations
Pooled Variance

Hypothesized Mean difference

193

df

Camphor

47.2371134

45.3125

79.93277491

81.48026316

97

96

80.70246802
0
191

t Stat

4.333985082

t Stat

5.440130482

t Stat

P(T<=t) one-tail

1.17452E-05

P(T<=t) one-tail

8.00339E-08

P(T<=t) one-tail

0.069181949

t CritiCamphorl one-tail

1.652747414

t CritiCamphorl one-tail

1.652788342

t CritiCamphorl one-tail

1.652870196

P(T<=t) two-tail

2.34904E-05

P(T<=t) two-tail

1.60068E-07

P(T<=t) two-tail

0.138363898

t CritiCamphorl two-tail

1.972266546

t CritiCamphorl two-tail

t CritiCamphorl two-tail

1.972462087

P(T<=t) two tail is

2.34904E-05

1.97233021

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

1.60068E-07

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference in rad length exists between Control and Cineole

* Sig difference in rad length exists btween Control and Camphor

* Mean rad length reduced in Cineole compared to Control

* Mean rad length reduced in Camphor compared to Control

1.48813704

P(T<=t) two tail is

0.138363898

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in rad length between Cineole and Camphor

Cupaniopsis parvifolia - Leaf Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

2.101010101

2.010309278

Mean

2.101010101

2.09375

Variance

0.459080602

0.322809278

Variance

0.459080602

0.275328947

99

97

99

96

Observations
Pooled Variance

0.39164737

Hypothesized Mean difference

df

Observations
Pooled Variance

0.368632896

Hypothesized Mean difference

194

2.010309278

2.09375

Variance

0.322809278

0.275328947

97

96

Observations
Pooled Variance

df

1.014469625

t Stat

P(T<=t) one-tail

0.155811128

P(T<=t) one-tail

t CritiCamphorl one-tail

1.652747414

t CritiCamphorl one-tail

P(T<=t) two-tail

0.311622256

P(T<=t) two-tail

0.93355654

P(T<=t) two-tail

t CritiCamphorl two-tail

1.972266546

t CritiCamphorl two-tail

1.97233021

t CritiCamphorl two-tail

0.311622256

0.083479824

P(T<=t) one-tail

1.652788342

t CritiCamphorl one-tail

0.93355654

value > 0.05 - null hypothesis (Ho) accepted

191
-1.059607128
0.14533075
1.652870196
0.2906615
1.972462087

P(T<=t) two tail is

0.2906615

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in leaf no. between Control and Camphor

* No sig dif exists in leaf no. between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-29

* No sig dif exists in leaf no. between Control and Cineole

t Stat

0.46677827

P(T<=t) two tail is

df

t Stat

P(T<=t) two tail is

0.299193407

Hypothesized Mean difference

193

Camphor

Mean

Cupaniopsis parvifolia - Leaf Area


t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

181.7777778

165.0412371

Mean

181.7777778

167.4583333

Mean

165.0412371

167.4583333

Variance

4354.664399

3426.706615

Variance

4354.664399

2942.019298

Variance

3426.706615

2942.019298

99

97

99

96

97

96

Observations
Pooled Variance

3895.468795

Hypothesized Mean difference


df

Observations
Pooled Variance

Hypothesized Mean difference

194

df

3659.320956

Observations
Pooled Variance

Hypothesized Mean difference

193

df

3185.631772
0
191

t Stat

1.876986984

t Stat

1.652577462

t Stat

P(T<=t) one-tail

0.031010508

P(T<=t) one-tail

0.050021367

P(T<=t) one-tail

0.383216594

t CritiCamphorl one-tail

1.652747414

t CritiCamphorl one-tail

1.652788342

t CritiCamphorl one-tail

1.652870196

P(T<=t) two-tail

0.062021015

P(T<=t) two-tail

0.100042735

P(T<=t) two-tail

0.766433187

t CritiCamphorl two-tail

1.972266546

t CritiCamphorl two-tail

t CritiCamphorl two-tail

1.972462087

P(T<=t) two tail is

0.062021015

1.97233021

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

*No sig dif exists in leaf area between Control and Cineole

0.100042735

value > 0.05 - null hypothesis (Ho) accepted

*No sig dif exists in leaf area between Control and Camphor

-0.297467129

P(T<=t) two tail is

0.766433187

value > 0.05 - null hypothesis (Ho) accepted

*No sig dif exists in leaf area between Cineole and Camphor

Cyperus enervis - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

29.09090909

30.41666667

Mean

29.09090909

20.16666667

Mean

30.41666667

20.16666667

Variance

194.0909091

170.2651515

Variance

194.0909091

20.92753623

Variance

170.2651515

20.92753623

11

12

11

24

12

24

Observations
Pooled Variance

181.6107504

Hypothesized Mean difference

t Stat

Pooled Variance

df

Observations

73.40128558

Hypothesized Mean difference

21
-0.235676559

Pooled Variance

df

Observations

69.24264706

Hypothesized Mean difference

33

df

34

t Stat

2.860799804

t Stat

3.484031139

P(T<=t) one-tail

0.407983433

P(T<=t) one-tail

0.003638773

P(T<=t) one-tail

0.000690081

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

1.692360456

t CritiCamphorl one-tail

1.690923455

P(T<=t) two-tail

0.815966866

P(T<=t) two-tail

0.007277546

P(T<=t) two-tail

0.001380161

t CritiCamphorl two-tail

2.079614205

t CritiCamphorl two-tail

t CritiCamphorl two-tail

2.032243174

P(T<=t) two tail is

0.815966866

2.03451691

P(T<=t) two tail is

0.007277546

value > 0.05 - null hypothesis (Ho) accepted

0.001380161

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Camphor

*Sig difference exists in shoot length between Cineole and Camphor

* Mean shoot length sig decreased in Camphor below Control

* Mean shoot length sig decreased in Camphor below Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-30

* No Sig difference exists in shoot length between Control and Cineole

P(T<=t) two tail is

Cyperus enervis - Radical


Length
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

21.54545455

23.75

Variance

111.2727273

164.2045455

11

12

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

138.9989177
0

13.75

Variance

111.2727273

105.673913

11

24

Observations

Hypothesized Mean difference


df

107.3705234

Mean
Variance
Observations
Pooled Variance

Hypothesized Mean difference

33

df

Camphor

23.75

13.75

164.2045455

105.673913

12

24

124.6102941
0
34

t Stat

2.066173798

t Stat

2.533774919

P(T<=t) one-tail

0.329385688

P(T<=t) one-tail

0.023367353

P(T<=t) one-tail

0.008030409

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

1.692360456

t CritiCamphorl one-tail

1.690923455

P(T<=t) two-tail

0.658771376

P(T<=t) two-tail

0.046734707

P(T<=t) two-tail

0.016060817

t CritiCamphorl two-tail

2.079614205

t CritiCamphorl two-tail

t CritiCamphorl two-tail

2.032243174

P(T<=t) two tail is

-0.447957071

Cineole

21.54545455

Pooled Variance

21

Camphor

Mean

0.658771376

2.03451691

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Cineole

0.046734707

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.016060817

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Camphor

*Sig difference exists in rad. length between Cineole and Camphor

* Mean rad length sig decreased in Camphor below Control

* Mean rad length sig decreased in Camphor below Cineole

Cyperus enervis - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

3.636363636

3.083333333

Mean

3.636363636

2.833333333

Mean

3.083333333

2.833333333

Variance

0.854545455

2.083333333

Variance

0.854545455

0.405797101

Variance

2.083333333

0.405797101

11

12

11

24

12

24

Observations
Pooled Variance

1.498196248

Hypothesized Mean difference

Pooled Variance

df

Observations

0.541781451

Hypothesized Mean difference

21

Pooled Variance

df

Observations

0.948529412

Hypothesized Mean difference

33

df

34

t Stat

1.082399602

t Stat

2.996317352

t Stat

0.726038417

P(T<=t) one-tail

0.145673879

P(T<=t) one-tail

0.002577835

P(T<=t) one-tail

0.236391951

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

1.692360456

t CritiCamphorl one-tail

1.690923455

P(T<=t) two-tail

0.291347758

P(T<=t) two-tail

0.00515567

P(T<=t) two-tail

0.472783902

t CritiCamphorl two-tail

2.079614205

t CritiCamphorl two-tail

2.03451691

t CritiCamphorl two-tail

2.032243174

P(T<=t) two tail is

0.291347758

P(T<=t) two tail is

0.00515567

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf No. between Control and Cineole

P(T<=t) two tail is

0.472783902

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf No. between Control and Camphor

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf No. between Cineole and Camphor

A11-31

* Mean leaf no. sig reduced in Camphor below Control

Cyperus enervis - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

14.54545455

15.20833333

Mean

14.54545455

10.08333333

Mean

15.20833333

10.08333333

Variance

48.52272727

42.56628788

Variance

48.52272727

5.231884058

Variance

42.56628788

5.231884058

11

12

11

24

12

24

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

45.40268759

Observations
Pooled Variance

Hypothesized Mean difference

21

df

-0.235676559

18.3503214

Observations
Pooled Variance

Hypothesized Mean difference

33

df

17.31066176
0
34

t Stat

2.860799804

t Stat

3.484031139

P(T<=t) one-tail

0.407983433

P(T<=t) one-tail

0.003638773

P(T<=t) one-tail

0.000690081

t CritiCamphorl one-tail

1.720743512

t CritiCamphorl one-tail

1.692360456

t CritiCamphorl one-tail

1.690923455

P(T<=t) two-tail

0.815966866

P(T<=t) two-tail

0.007277546

P(T<=t) two-tail

0.001380161

t CritiCamphorl two-tail

2.079614205

t CritiCamphorl two-tail

t CritiCamphorl two-tail

2.032243174

P(T<=t) two tail is

0.815966866

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Cineole

2.03451691

P(T<=t) two tail is

0.007277546

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.001380161

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Control and Camphor

* Sig difference exists in leaf area between Cineole and Camphor

* Mean leaf area sig reduced in Camphor compared to Control

* Mean leaf area sig. reduced in Camphor below Cineole

Eucalyptus intermedia - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

75.25806452

57.5

Variance

142.1500701

154.2977528

93

90

Observations
Pooled Variance

148.1232401

Hypothesized Mean difference

t Stat

1.653315849

P(T<=t) two-tail

1.19488E-18

t CritiCamphorl two-tail

1.97315785

P(T<=t) two tail is

Mean

Variance

142.1500701

114.5196353

Variance

93

93

Observations

t Stat

5.97E-19

t CritiCamphorl one-tail

75.74193548

128.3348527

1.19488E-18

75.74193548

154.2977528

114.5196353

90

93

134.0790412

Hypothesized Mean difference

184

df

-0.291261937

t Stat

Camphor
57.5

Observations
Pooled Variance

df

9.867819843

Cineole

75.25806452

Hypothesized Mean difference

181

P(T<=t) one-tail

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

181
-10.65436698

P(T<=t) one-tail

0.385589535

P(T<=t) one-tail

3.47064E-21

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653315849
6.94128E-21

P(T<=t) two-tail

0.771179069

P(T<=t) two-tail

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

P(T<=t) two tail is

0.771179069

value < 0.05 - null hypothesis (Ho) rejected

1.97315785

P(T<=t) two tail is

6.94128E-21

value > 0.05 - null hypothesis (Ho) is not rejected

value < 0.05 - null hypothesis (Ho) is rejected

* no significant difference exists in shoot length between the Control and Camphor treatments.

* significant difference in shoot length between Cineole and Camphor treatments.

* Mean length > in the Control than Cineole indicating reduced shoot growth Cineole treatment

* Mean lengths Control and Camphor similar - no significant effect of Camphor on shoot length.

* Mean length reduced in Cineole treatment below Camphor.

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-32

* A significant difference exists in shoot length between the Control and Cineole treatments.

Length
Eucalyptus intermedia - Radical
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

56.03225806

56.46666667

Mean

56.03225806

54.80645161

Mean

56.46666667

54.80645161

Variance

256.4880785

275.0606742

Variance

256.4880785

143.0273492

Variance

275.0606742

143.0273492

93

90

93

93

90

93

Observations
Pooled Variance

265.6204598

Hypothesized Mean difference

t Stat

Pooled Variance

df

Observations

199.7577139

Hypothesized Mean difference

181

df

-0.180262271

Observations
Pooled Variance

Hypothesized Mean difference

184

df

207.9498129
0
181

t Stat

0.591420807

t Stat

0.778614101

P(T<=t) one-tail

0.277482193

P(T<=t) one-tail

0.218611888

P(T<=t) one-tail

0.428574169

t CritiCamphorl one-tail

1.653315849

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653315849

P(T<=t) two-tail

0.857148338

P(T<=t) two-tail

0.554964386

P(T<=t) two-tail

0.437223777

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

t CritiCamphorl two-tail

1.97315785

P(T<=t) two tail is

0.857148338

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) not rejected

*no significant difference between Control and Cineole

0.554964386

1.97315785

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) not rejected

*no significant difference between Control and Camphor

0.437223777

value > 0.05 - null hypothesis (Ho) not rejected

*no significant difference between Cineolel and Camphor

Eucalyptus intermedia - Leaf Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean

Control

4.139784946

3.233333333

Mean

0.77372604

1.282022472

Variance

93

90

Variance
Observations
Pooled Variance

Cineole

1.023661855

Hypothesized Mean difference

t Stat

Camphor

Mean

3.233333333

4.075268817

0.77372604

1.048620851

Variance

1.282022472

1.048620851

93

93

90

93

0.911173446

1.163387394

Hypothesized Mean difference

184

t Stat

Observations
Pooled Variance

df

6.05903624

Cineole

4.075268817

Hypothesized Mean difference

181

Camphor

4.139784946

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

df

0.46088678

t Stat

181
-5.279026242

P(T<=t) one-tail

3.87767E-09

P(T<=t) one-tail

0.322712062

P(T<=t) one-tail

1.84537E-07

t CritiCamphorl one-tail

1.653315849

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653315849

P(T<=t) two-tail

7.75535E-09

P(T<=t) two-tail

0.645424124

P(T<=t) two-tail

3.69074E-07

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

t CritiCamphorl two-tail

1.97315785

P(T<=t) two tail is

7.75535E-09

P(T<=t) two tail is

0.645424124

value < 0.05 - null hypothesis (Ho) rejected

1.97315785

P(T<=t) two tail is

3.69074E-07

value > 0.05 - null hypothesis (Ho) not rejected

value < 0.05 - null hypothesis (Ho) rejected

* No significant difference in leaf No. exists between the Control and Camphor treatments.

* Their is a significant difference in Leaf No. between the Cineole and Camphor treatments.

* Mean length > in Control than Cineole - sig. reduced leaf production Cineole treatment

* Mean length similar in Control and Camphor - no effect on leaf No. due to Camphor treatment

* Mean Leaf No. is reduced in the Cineole compared to Camphor treatment.

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-33

* A significant difference exists in leaf No. exists between the Control and Cineole treatments.

Eucalyptus intermedia - Leaf Area


t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean difference
df

Cineole

Control

121.516129

79.5

2716.774194

1332.185393

93

90

2035.954286

Mean
Variance
Observations
Pooled Variance

Hypothesized Mean difference

181

df

Camphor

Cineole

121.516129

79.58064516

2716.774194

1169.81136

93

93

1943.292777
0
184

Variance
Observations
Pooled Variance
Hypothesized Mean difference
df

79.58064516

1332.185393

1169.81136

90

93

1249.652736
0
181

6.297515006

t Stat

P(T<=t) one-tail

1.11451E-09

P(T<=t) one-tail

3.93017E-10

P(T<=t) one-tail

0.493853701

t CritiCamphorl one-tail

1.653315849

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653315849

P(T<=t) two-tail

2.22903E-09

P(T<=t) two-tail

7.86033E-10

P(T<=t) two-tail

0.987707401

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

P(T<=t) two tail is

7.86033E-10

P(T<=t) two tail is

1.97315785

2.22903E-09

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

t Stat

Camphor
79.5

t Stat

t CritiCamphorl two-tail

6.48692709

Mean

-0.015428398

1.97315785

0.987707401

value > 0.05 - null hypothesis (Ho) not rejected

* A significant difference exists in leaf Area between the Control and Cineole treatments.

* A significant difference in leal Area exists between the Control and Camphor treatments.

* Their is a significant difference in Leaf Are. between the Cineole and Camphor treatments.

* Mean length > in Control than Cineole - significantly reduced leaf area Cineole treatment

* Mean Area is significantly reduced in the Camphor treatment indicating effect.

* Mean Leaf No. is similar indicating not difference in effect.

Eucalyptus saligna - Shoot Length


t-Test: Two-Sample Assuming Equal Variances
t-Test: Two-Sample Assuming Equal Variances

Mean

21.63636364

18.265625

Variance

48.88111888

25.84895833

66

64

Observations
Pooled Variance

37.54497736

Hypothesized Mean difference

128

Cineole

19.58441558

Mean

Variance

48.88111888

32.35133288

Variance

66

77

39.97144699

Hypothesized Mean difference

t Stat

1.934820759

t Stat

1.656844688

P(T<=t) one-tail

0.027507208

P(T<=t) one-tail

P(T<=t) two-tail

0.002126801

t CritiCamphorl one-tail

1.655732831

t CritiCamphorl one-tail

t CritiCamphorl two-tail

1.978669388

P(T<=t) two-tail

0.055014415

P(T<=t) two-tail

t CritiCamphorl two-tail

1.976932253

t CritiCamphorl two-tail

0.002126801

P(T<=t) two tail is

0.055014415

value < 0.05 - null hypothesis (Ho) rejected

32.35133288

64

77

29.40421348
0

df

t CritiCamphorl one-tail

P(T<=t) two tail is

19.58441558

25.84895833

Hypothesized Mean difference

141

Camphor

18.265625

Observations
Pooled Variance

df

0.0010634

Camphor

21.63636364

Pooled Variance

3.135735584

P(T<=t) one-tail

Control
Mean

Observations

df
t Stat

t-Test: Two-Sample Assuming Equal Variances

Cineole

139
-1.437794598
0.07636992
1.655889719
0.15273984
1.977177817

P(T<=t) two tail is

0.15273984

value > 0.05 - null hypothesis (Ho) not rejected

value > 0.05 - null hypothesis (Ho) not rejected

* A significant difference exists in shoot length between the Control and Cineole treatments.

* Their is no significant difference in shoot length between the Control and Camphor treatments.

* Their is no significant difference in shoot length between the Cineole and Camphor treatments.

* Mean length is significantly reduced in the Cineole treatment

* Mean length of both treatments similar.

* Mean shoot length is similar in both treatments.

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Length
Eucalyptus saligna - Radical
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

20.34848485

22.453125

Variance

60.78438228

118.3152282

66

64

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

89.1003455
0

17.62337662

Mean

Variance

60.78438228

52.68523582

Variance

66

77

Observations

Hypothesized Mean difference


df

-1.270949914

Cineole

20.34848485

Pooled Variance

128

Camphor

Mean

t Stat

56.4188849

Observations
Pooled Variance

Hypothesized Mean difference

141

df

2.162822192

Camphor

22.453125

17.62337662

118.3152282

52.68523582

64

77

82.43120358
0
139

t Stat

3.144882892

P(T<=t) one-tail

0.001016132

P(T<=t) one-tail

0.103025697

P(T<=t) one-tail

t CritiCamphorl one-tail

1.656844688

t CritiCamphorl one-tail

1.655732831

t CritiCamphorl one-tail

1.655889719

P(T<=t) two-tail

0.206051393

P(T<=t) two-tail

0.032241039

P(T<=t) two-tail

0.002032263

t CritiCamphorl two-tail

1.978669388

t CritiCamphorl two-tail

1.976932253

t CritiCamphorl two-tail

1.977177817

P(T<=t) two tail is

0.206051393

value > 0.05 - null hypothesis (Ho) not rejected

*No significant effect on rad. length can be attributed to the Cineole treatment.

P(T<=t) two tail is

0.01612052

0.032241039

value < 0.05 - null hypothesis (Ho) rejected

* Radical length was significantly decreased in the Camphor treatment compared to the Control

P(T<=t) two tail is

A11-34

Control

0.002032263

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference radical length between Cineole and Camphor treatments - Camphor sig. reduced.

Eucalyptus saligna - Leaf Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

4.590909091

3.96875

Variance

2.245454545

2.094246032

66

64

Observations
Pooled Variance

2.171031605

Hypothesized Mean difference

Cineole

4.590909091

2.831168831

Mean

Variance

2.245454545

1.615857826

Variance

66

77

Observations

1.906097449

Hypothesized Mean difference

128

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

df

2.831168831

2.094246032

1.615857826

64

77

Observations
Pooled Variance

1.832681258

Hypothesized Mean difference

141

Camphor

3.96875

df

139

t Stat

2.406903381

t Stat

7.598455627

t Stat

P(T<=t) one-tail

0.008758076

P(T<=t) one-tail

1.89752E-12

P(T<=t) one-tail

4.967807037
9.78471E-07

t CritiCamphorl one-tail

1.656844688

t CritiCamphorl one-tail

1.655732831

t CritiCamphorl one-tail

1.655889719

P(T<=t) two-tail

0.017516152

P(T<=t) two-tail

3.79503E-12

P(T<=t) two-tail

1.95694E-06

t CritiCamphorl two-tail

1.978669388

t CritiCamphorl two-tail

1.976932253

t CritiCamphorl two-tail

1.977177817

P(T<=t) two tail is

0.017516152

P(T<=t) two tail is

3.79503E-12

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

1.95694E-06

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

*A significant difference in leaf No. exists between the Control and Camphor treatments.

* A significant difference in Leaf No. exists between the Cineole and Camphor treatments.

* Mean leaf No. is reduced in the Cineole treatment

* Mean length is reduced in the Camphor treatment

* Mean leaf number reduced in Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-35

* A significant difference in leaf No. exists between the Control and Cineole treatments.

Eucalyptus saligna - Leaf Area


t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

12.96923077

8.671875

Variance

152.8427885

57.36681548

65

64

Observations
Pooled Variance

105.4806916

Hypothesized Mean difference

t Stat

4.077922078

Mean

Variance

152.8427885

12.78332194

Variance

65

77

Observations

Hypothesized Mean difference

127

df

2.37611051

Cineole

12.96923077

Pooled Variance

df

Camphor

Mean

76.81050664
0

4.077922078

57.36681548

12.78332194

64

77

Observations
Pooled Variance

32.99022908

Hypothesized Mean difference

140

Camphor

8.671875

df

139

t Stat

6.023005259

t Stat

P(T<=t) one-tail

0.009495317

P(T<=t) one-tail

7.15468E-09

P(T<=t) one-tail

4.728459438
2.74417E-06

t CritiCamphorl one-tail

1.656940185

t CritiCamphorl one-tail

1.655810138

t CritiCamphorl one-tail

1.655889719

P(T<=t) two-tail

0.018990634

P(T<=t) two-tail

1.43094E-08

P(T<=t) two-tail

5.48833E-06

t CritiCamphorl two-tail

1.978819455

t CritiCamphorl two-tail

1.977055035

t CritiCamphorl two-tail

1.977177817

P(T<=t) two tail is

0.018990634

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

1.43094E-08

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

5.48833E-06

value < 0.05 - null hypothesis (Ho) rejected

* a significant difference exists in leaf Area between the Control and Cineole treatments.

* A significant difference in leal Area exists between the Control and Camphor treatments.

* A significant difference in leaf Area exists between the Cineole and Camphor treatments.

* Mean length is reduced in the Cineole treatment

* Mean leaf Area is significantly reduced in the Camphor treatment indicating effect.

* Mean Area is reduced in the Camphor treatment.

Ficus coronata - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

6.133333333

13.20652174

Mean

6.133333333

10.6

Variance

3.838095238

14.71512183

Variance

3.838095238

14.42025316

15

92

15

80

Observations
Pooled Variance

13.26485162

Hypothesized Mean difference

Pooled Variance

df

Observations

12.82724014

Hypothesized Mean difference

105

13.20652174

10.6

Variance

14.71512183

14.42025316

92

80

Observations
Pooled Variance

df

14.57809463

Hypothesized Mean difference

93

df

-6.97447758

t Stat

P(T<=t) one-tail

1.41621E-10

P(T<=t) one-tail

t CritiCamphorl one-tail

1.659495865

t CritiCamphorl one-tail

P(T<=t) two-tail

2.83242E-10

P(T<=t) two-tail

2.54564E-05

P(T<=t) two-tail

1.45038E-05

t CritiCamphorl two-tail

1.982816684

t CritiCamphorl two-tail

1.985799827

t CritiCamphorl two-tail

1.974017323

2.83242E-10

t Stat

170

t Stat

P(T<=t) two tail is

-4.432469453

Camphor

Mean

1.27282E-05
1.66140353

P(T<=t) two tail is

2.54564E-05

value < 0.05 - null hypothesis (Ho) rejected

4.465659983

P(T<=t) one-tail

7.25191E-06

t CritiCamphorl one-tail

1.653866093

P(T<=t) two tail is

1.45038E-05

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Camphor

*Sig difference exists in shoot length between Cineole and Camphor

* Mean shoot length sig increased in Cineole above Control

* Mean shoot length sig increased in Camphor above Control

* Mean shoot length sig greater in Cineole than Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-36

* Sig difference exists in shoot length between Control and Cineole

Ficus coronata - Radical


Length
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

2.466666667

7.293478261

Variance

0.980952381

16.7810559

15

92

Observations
Pooled Variance

14.67437543

Hypothesized Mean difference

t Stat

5.35

Variance

0.980952381

6.660759494

15

80

Observations

5.805734767

Hypothesized Mean difference

105

t Stat

P(T<=t) one-tail

7.98073E-06

P(T<=t) one-tail

t CritiCamphorl one-tail

1.659495865

t CritiCamphorl one-tail

P(T<=t) two-tail

1.59615E-05

P(T<=t) two-tail

t CritiCamphorl two-tail

1.982816684

t CritiCamphorl two-tail

P(T<=t) two tail is

1.59615E-05

Mean
Variance

5.35

16.7810559

6.660759494

92

80

12.07809463

Hypothesized Mean difference

93

df

-4.252997011
2.50948E-05
1.66140353

Camphor

7.293478261

Observations
Pooled Variance

df

-4.525100079

Cineole

2.466666667

Pooled Variance

df

Camphor

Mean

170

t Stat

3.658096491

P(T<=t) one-tail

0.000169326

t CritiCamphorl one-tail

1.653866093

5.01896E-05

P(T<=t) two-tail

0.000338652

1.985799827

t CritiCamphorl two-tail

1.974017323

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

5.01896E-05

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejectted

0.000338652

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Cineole

* Sig difference exists in rad. length between Control and Camphor

*Sig difference exists in rad. length between Cineole and Camphor

* Mean rad length increased in Cineole above Control

* Mean rad length increased in Camphor above Control

* Mean rad length greater in Cineole than Camphor

Ficus coronata - Leaf Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

1.133333333

0.943820225

Mean

1.133333333

0.4125

Variance

0.266666667

0.417262513

Variance

0.266666667

0.245411392

15

89

15

80

Observations
Pooled Variance

0.396592495

Hypothesized Mean difference

Pooled Variance

df

Observations

0.248611111

Hypothesized Mean difference

102

0.943820225

0.4125

Variance

0.417262513

0.245411392

89

80

Observations
Pooled Variance

df

0.335967671

Hypothesized Mean difference

93

Camphor

Mean

df

167

t Stat

1.078179259

t Stat

5.138110199

t Stat

P(T<=t) one-tail

0.141748478

P(T<=t) one-tail

7.62206E-07

P(T<=t) one-tail

t CritiCamphorl one-tail

1.659930149

t CritiCamphorl one-tail

P(T<=t) two-tail

0.283496956

P(T<=t) two-tail

1.52441E-06

P(T<=t) two-tail

1.53222E-08

t CritiCamphorl two-tail

1.983494258

t CritiCamphorl two-tail

1.985799827

t CritiCamphorl two-tail

1.974271981

P(T<=t) two tail is

0.283496956

1.66140353

P(T<=t) two tail is

1.52441E-06

value > 0.05 - null hypothesis (Ho) accepted

7.6611E-09
1.654029802

P(T<=t) two tail is

1.53222E-08

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf No. between Control and Camphor

* Sig difference exists in leaf No. between Cineole and Camphor

* Mean leaf no. sig reduced in Camphor below Control

* Mean leaf no. reduced in Camphor below Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-37

* No sig difference exists in leaf No. between Control and Cineole

t CritiCamphorl one-tail

5.949825758

Ficus coronata - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

1.714285714

3.402173913

Mean

1.714285714

0.518987342

Mean

3.402173913

0.518987342

Variance

1.296703297

14.63867654

Variance

1.296703297

0.611814346

Variance

14.63867654

0.611814346

14

92

14

79

92

79

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

12.97092989

Observations
Pooled Variance

0.709655625

Hypothesized Mean difference

104

8.164740143

Hypothesized Mean difference

91

df

169

t Stat

4.893150715

t Stat

P(T<=t) one-tail

0.052676465

P(T<=t) one-tail

2.13324E-06

P(T<=t) one-tail

2.85747E-10

t CritiCamphorl one-tail

1.659636837

t CritiCamphorl one-tail

1.661771876

t CritiCamphorl one-tail

1.653920663

P(T<=t) two-tail

4.26647E-06

P(T<=t) two-tail

5.71495E-10

t CritiCamphorl two-tail

1.986377356

t CritiCamphorl two-tail

1.974099177

P(T<=t) two-tail
t CritiCamphorl two-tail

-1.633665433

Pooled Variance

df

Observations

0.10535293
1.983034963

P(T<=t) two tail is

0.10535293

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in leaf area between Control and Cineole

4.26647E-06

6.578258977

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

5.71495E-10

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Control and Camphor

* Sig difference exists in leaf area between Cineole and Camphor

* Mean leaf area reduced in Camphor below Control

* Mean leaf area greater in Cineole than Camphor

Guioa semiglauca - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

67.56060606

59

Variance

59.63473193

86.69333333

66

76

Observations
Pooled Variance

74.13041126

Hypothesized Mean difference

Cineole

67.56060606

66.94186047

Mean

Variance

59.63473193

101.4201094

Variance

66

86

Observations

83.31311252

Hypothesized Mean difference

140

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

df

66.94186047

86.69333333

101.4201094

76

86

Observations
Pooled Variance

94.51693314

Hypothesized Mean difference

150

Camphor
59

df

160

t Stat

5.909366138

t Stat

0.414242677

t Stat

-5.18878835

P(T<=t) one-tail

1.24605E-08

P(T<=t) one-tail

0.339644071

P(T<=t) one-tail

3.16754E-07

t CritiCamphorl one-tail

1.655810138

t CritiCamphorl one-tail

1.655075721

t CritiCamphorl one-tail

1.654432253

P(T<=t) two-tail

2.49211E-08

P(T<=t) two-tail

0.679288143

P(T<=t) two-tail

6.33509E-07

t CritiCamphorl two-tail

1.977055035

t CritiCamphorl two-tail

1.975904524

t CritiCamphorl two-tail

P(T<=t) two tail is

2.49211E-08

P(T<=t) two tail is

0.679288143

value < 0.05 - null hypothesis (Ho) rejected

1.97490408

P(T<=t) two tail is

6.33509E-07

value > 0.05 - null hypothesis (Ho) accepted

value < 0.05 - null hypothesis (Ho) rejected

*No sig.dif in shoot length between Control and Camphor treatments

*Sig dif in shoot length between Cineole and Camphor treatments

* Mean shhot length similar

* Mean shoot length increased in Camphor treatment

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-38

*Sig.dif in shoot length exists between Control and Cineole treatments


*Mean shoot length reduced in Cineole treatmemt

Radicle
Length
Guioa semiglauca - Radical
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

45.22727273

41.75324675

Mean

45.22727273

46.45348837

Mean

41.75324675

46.45348837

Variance

91.87062937

109.7146275

Variance

91.87062937

91.68604651

Variance

109.7146275

91.68604651

66

77

66

86

77

86

Observations
Pooled Variance

101.4886709

Hypothesized Mean difference

t Stat

Pooled Variance

df

Observations

Hypothesized Mean difference

141

df

2.05576558

t Stat

91.76603242

Observations
Pooled Variance

Hypothesized Mean difference

150

df

-0.782213442

t Stat

100.1964326
0
161
-2.992918152

P(T<=t) one-tail

0.020824993

P(T<=t) one-tail

0.217661093

P(T<=t) one-tail

0.001599506

t CritiCamphorl one-tail

1.655732831

t CritiCamphorl one-tail

1.655075721

t CritiCamphorl one-tail

1.654373136

P(T<=t) two-tail

0.041649987

P(T<=t) two-tail

0.435322187

P(T<=t) two-tail

0.003199012

t CritiCamphorl two-tail

1.976932253

t CritiCamphorl two-tail

1.975904524

t CritiCamphorl two-tail

1.974808583

P(T<=t) two tail is

0.041649987

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.435322187

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

0.003199012

value < 0.05 - null hypothesis (Ho) rejected

* No sig. dif in rad. length between Control and Camphor treatments


* Sig.dif exists in rad. length between Control and Cineole treatments
* Mean rad. length is reduced in the Cineole treatment

* Mean rad length similar

* sg. dif in rad length exist between Cineole and Camphor treatments
*Cineole has reduced mean rad. length

Guioa semiglauca - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

2.03030303

2.012987013

Mean

2.03030303 1.942528736

Mean

2.012987013

1.942528736

Variance

0.02983683

0.012987013

Variance

0.02983683 0.124565624

Variance

0.012987013

0.124565624

66

77

77

87

Observations
Pooled Variance

0.020754659

Hypothesized Mean difference

Pooled Variance

df

Observations

66
0.083788329

Hypothesized Mean difference

141

87

Pooled Variance

df

Observations

0.072220103

Hypothesized Mean difference

151

df

162

t Stat

0.716538485

t Stat

1.857639612

t Stat

1.675660508

P(T<=t) one-tail

0.237422035

P(T<=t) one-tail

0.032583852

P(T<=t) one-tail

0.047866865

t CritiCamphorl one-tail

1.655732831

t CritiCamphorl one-tail

1.655007509

t CritiCamphorl one-tail

1.654314019

P(T<=t) two-tail

0.065167704

P(T<=t) two-tail

t CritiCamphorl two-tail

1.975799933

t CritiCamphorl two-tail

P(T<=t) two-tail
t CritiCamphorl two-tail

0.47484407
1.976932253

P(T<=t) two tail is

0.47484407

P(T<=t) two tail is

0.065167704

value > 0.05 - null hypothesis (Ho) accepted

P(T<=t) two tail is

0.09573373

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

*No sig. difference in leaf No.for Camphor

*No Sig. difference in leaf No.between Cineole and Camphor.

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-39

*No sig. difference in leaf No.for Cineole

0.09573373
1.974717634

Guioa semiglauca - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

59.34848485

44.87012987

Mean

59.34848485 34.97674419

Mean

44.87012987

34.97674419

Variance

613.2151515

429.8513329

Variance

613.2151515 416.6582763

Variance

429.8513329

416.6582763

66

77

77

86

Observations
Pooled Variance
Hypothesized Mean difference
df

514.3807528

Observations
Pooled Variance

Hypothesized Mean difference

141

df

66

86

501.8329222

Observations
Pooled Variance

Hypothesized Mean difference

150

df

422.8860546
0
161

t Stat

3.805626628

t Stat

6.648235162

t Stat

3.066436983

P(T<=t) one-tail

0.000105085

P(T<=t) one-tail

2.58618E-10

P(T<=t) one-tail

0.001270739

t CritiCamphorl one-tail

1.655732831

t CritiCamphorl one-tail

1.655075721

t CritiCamphorl one-tail

1.654373136

P(T<=t) two-tail

5.17237E-10

P(T<=t) two-tail

0.002541478

t CritiCamphorl two-tail

1.975904524

t CritiCamphorl two-tail

1.974808583

P(T<=t) two-tail
t CritiCamphorl two-tail

0.00021017
1.976932253

P(T<=t) two tail is

0.00021017

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

5.17237E-10

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.002541478

value < 0.05 - null hypothesis (Ho) rejected

*Sig.difference in leaf area exists between Control and Cineole

*Sig.difference in leaf area exists between Control and Camphor

*Sig.difference in leaf area exists between Cineole and Camphor

*Mean leaf area decreased in Cineole.

*Mean leaf area decreased in Camphor.

*Mean leaf area decreased in Camphor.

Helichrysum bracteatum - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Mean
Variance

Pooled Variance

7.333333333

Mean

9.333333333

5.066666667

Variance

7.393939394

Hypothesized Mean difference

0.14705686

Camphor

Mean

7.333333333

7.153846154

9.333333333

5.641025641

Variance

5.066666667

5.641025641

13

13

6.871794872

Observations
Pooled Variance

df

1.101701081

P(T<=t) one-tail

Cineole

7.153846154

Hypothesized Mean difference

11

Camphor
9

Observations
Pooled Variance

df
t Stat

Control

Observations

t-Test: Two-Sample Assuming Equal Variances

5.472096531

Hypothesized Mean difference

18

df

t Stat

1.502237138

t Stat

P(T<=t) one-tail

0.075188187

P(T<=t) one-tail

17
0.155463005
0.43914367

t CritiCamphorl one-tail

1.795883691

t CritiCamphorl one-tail

1.734063062

t CritiCamphorl one-tail

1.739606432

P(T<=t) two-tail

0.294113719

P(T<=t) two-tail

0.150376373

P(T<=t) two-tail

0.878287341

t CritiCamphorl two-tail

2.200986273

t CritiCamphorl two-tail

2.100923666

t CritiCamphorl two-tail

2.109818524

P(T<=t) two tail is

0.294113719

P(T<=t) two tail is

0.150376373

value > 0.05 - null hypothesis (Ho) accepted

0.878287341

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in shoot length between Control and Camphor

*No Sig difference exists in shoot length between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-40

* No Sig difference exists in shoot length between Control and Cineole

P(T<=t) two tail is

Length
Helichrysum bracteatum - Radical
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

27.14285714

18.83333333

Mean

27.14285714

14.76923077

Mean

18.83333333

14.76923077

Variance

293.4761905

166.1666667

Variance

293.4761905

63.02564103

Variance

166.1666667

63.02564103

13

13

Observations
Pooled Variance
Hypothesized Mean difference
df

235.6082251

Observations
Pooled Variance

Hypothesized Mean difference

11

df

139.8424908

Observations
Pooled Variance

Hypothesized Mean difference

18

df

93.3612368
0
17

t Stat

0.973047143

t Stat

2.231943112

t Stat

0.852220275

P(T<=t) one-tail

0.175720802

P(T<=t) one-tail

0.019283523

P(T<=t) one-tail

0.202969136

t CritiCamphorl one-tail

1.795883691

t CritiCamphorl one-tail

1.734063062

t CritiCamphorl one-tail

1.739606432

P(T<=t) two-tail

0.351441604

P(T<=t) two-tail

0.038567046

P(T<=t) two-tail

0.405938272

t CritiCamphorl two-tail

2.200986273

t CritiCamphorl two-tail

2.100923666

t CritiCamphorl two-tail

2.109818524

P(T<=t) two tail is

0.351441604

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Cineole

0.038567046

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Camphor


* Mean rad length sig decreased in Camphor below Control

P(T<=t) two tail is

0.405938272

value > 0.05 - null hypothesis (Ho) accepted

*No Sig difference exists in rad. length between Cineole and Camphor

Helichrysum bracteatum - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

1.333333333

0.8

1.090909091

Hypothesized Mean difference

Mean
Variance

Mean

1.333333333

1.397435897

Variance

13

1.376068376

t Stat

Observations
Pooled Variance

df

-1.720912102

Cineole

4.307692308

Hypothesized Mean difference

11

Camphor
4

Observations
Pooled Variance

df
t Stat

Control

Observations
Pooled Variance

Cineole

t-Test: Two-Sample Assuming Equal Variances

1.397435897

13

df

17

t Stat

1.269065135

P(T<=t) one-tail

0.056617021

P(T<=t) one-tail

0.291357197

P(T<=t) one-tail

0.110759312

t CritiCamphorl one-tail

1.795883691

t CritiCamphorl one-tail

1.734063062

t CritiCamphorl one-tail

1.739606432

P(T<=t) two-tail

0.113234042

P(T<=t) two-tail

0.582714395

P(T<=t) two-tail

0.221518624

t CritiCamphorl two-tail

2.200986273

t CritiCamphorl two-tail

2.100923666

t CritiCamphorl two-tail

2.109818524

P(T<=t) two tail is

0.113234042

-0.559502885

4.307692308

0.8

1.221719457

Hypothesized Mean difference

18

Camphor
5

P(T<=t) two tail is

0.582714395

value > 0.05 - null hypothesis (Ho) accepted

0.221518624

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf No. between Control and Camphor

* No Sig difference exists in leaf No. between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-41

* No Sig difference exists in leaf No. between Control and Cineole

P(T<=t) two tail is

Helichrysum bracteatum - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean difference
df

Cineole

Control

19

11.66666667

Mean

23.33333333

57.86666667

Variance

39.03030303

Observations
Pooled Variance

Hypothesized Mean difference

11

df

Camphor

Cineole

Camphor

19

7.307692308

Mean

11.66666667

7.307692308

23.33333333

18.23076923

Variance

57.86666667

18.23076923

13

13

19.93162393

Observations
Pooled Variance

Hypothesized Mean difference

18

df

29.88838612
0
17

t Stat

2.109859004

t Stat

5.586432965

t Stat

P(T<=t) one-tail

0.029295582

P(T<=t) one-tail

1.32874E-05

P(T<=t) one-tail

t CritiCamphorl one-tail

1.795883691

t CritiCamphorl one-tail

1.734063062

t CritiCamphorl one-tail

1.739606432

P(T<=t) two-tail

0.058591164

P(T<=t) two-tail

2.65749E-05

P(T<=t) two-tail

0.124610501

t CritiCamphorl two-tail

2.200986273

t CritiCamphorl two-tail

2.100923666

t CritiCamphorl two-tail

2.109818524

P(T<=t) two tail is

0.058591164

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Cineole

P(T<=t) two tail is

2.65749E-05

value < 0.05 - null hypothesis (Ho) rejected

*Sig difference exists in leaf area between Control and Camphor


* Mean leaf area sig reduced in Camphor compared to Control

P(T<=t) two tail is

1.615487428
0.06230525

0.124610501

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Cineole and Camphor

Hymenosporum flavum - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

65.38461538

41.16

Mean

65.38461538

57.3030303

Mean

41.16

57.3030303

Variance

1118.589744

73.64

Variance

1118.589744

260.530303

Variance

73.64

260.530303

13

25

13

33

25

33

Observations
Pooled Variance

421.9565812

Hypothesized Mean difference

Pooled Variance

df

Observations

494.5465141

Hypothesized Mean difference

36

Pooled Variance

df

Observations

Hypothesized Mean difference

44

3.448838756

t Stat

1.109794855

t Stat

P(T<=t) one-tail

0.000725829

P(T<=t) one-tail

0.136558051

P(T<=t) one-tail

t CritiCamphorl one-tail

1.688297289

t CritiCamphorl one-tail

1.680230071

t CritiCamphorl one-tail

P(T<=t) two-tail

0.001451658

P(T<=t) two-tail

0.273116101

P(T<=t) two-tail

2.02809133

t CritiCamphorl two-tail

P(T<=t) two tail is

0.001451658

2.0153675

* Sig dif exists in shoot length between Control and Cineole

t CritiCamphorl two-tail

P(T<=t) two tail is

0.273116101

value < 0.05 - null hypothesis (Ho) rejected

df

t Stat

t CritiCamphorl two-tail

180.4344589

56
-4.532505072
1.5553E-05
1.672522103
3.1106E-05
2.003239388

P(T<=t) two tail is

3.1106E-05

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif in shoot length exists between Control and Camphor

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exist in shoot length exists between Cineole and Camphor
* mean shoot length reduced in Cineole below Camphor

A11-42

* Mean shoot length reduced in Cineole below Control

Radicle
Length
Length
Hymenosporum flavum - Radical
t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

42.30769231

51.6

Variance

440.0641026

632.75

13

25

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat
P(T<=t) one-tail

t-Test: Two-Sample Assuming Equal Variances

568.5213675
0

44.65116279

Mean

Variance

440.0641026

277.8516058

Variance

13

43

Observations

Hypothesized Mean difference


df

-1.139724983

t Stat

0.130963772

Cineole

42.30769231

Pooled Variance

36

Camphor

Mean

P(T<=t) one-tail

313.8988273

Observations
Pooled Variance

Hypothesized Mean difference

54

df

-0.417904003
0.338837229

Camphor
51.6

44.65116279

632.75

277.8516058

25

43

406.9055673
0
66

t Stat

1.369667332

P(T<=t) one-tail

0.087716475

t CritiCamphorl one-tail

1.688297289

t CritiCamphorl one-tail

1.673565748

t CritiCamphorl one-tail

1.668270215

P(T<=t) two-tail

0.261927544

P(T<=t) two-tail

0.677674458

P(T<=t) two-tail

0.175432949

t CritiCamphorl two-tail

2.004881026

t CritiCamphorl two-tail

1.996563697

t CritiCamphorl two-tail

2.02809133

P(T<=t) two tail is

0.261927544

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in rad length for Control and Cineole

P(T<=t) two tail is

0.677674458

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in rad length for Control and Camphor

P(T<=t) two tail is

0.175432949

value > 0.05 - null hypothesis (Ho) accepted

* No sig difference exists in rad length for Cineole and Camphor

Hymenosporum flavum - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

3.692307692

Variance

2.564102564

1.416666667

13

25

Observations
Pooled Variance

1.799145299

Hypothesized Mean difference

Cineole

3.692307692

2.837209302

Mean

Variance

2.564102564

1.282392027

Variance

13

43

Observations

1.56721659

Hypothesized Mean difference

36

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

2.837209302

1.416666667

1.282392027

25

43

Observations
Pooled Variance

df

Camphor
2

1.331219168

Hypothesized Mean difference

54

df

66

t Stat

3.689742779

t Stat

2.158059771

t Stat

P(T<=t) one-tail

0.000368741

P(T<=t) one-tail

0.017694007

P(T<=t) one-tail

t CritiCamphorl one-tail

1.688297289

t CritiCamphorl one-tail

1.673565748

t CritiCamphorl one-tail

1.668270215

P(T<=t) two-tail

0.000737482

P(T<=t) two-tail

0.035388014

P(T<=t) two-tail

0.005280985

t CritiCamphorl two-tail

2.004881026

t CritiCamphorl two-tail

1.996563697

t CritiCamphorl two-tail

2.02809133

P(T<=t) two tail is

0.000737482

P(T<=t) two tail is

0.035388014

value < 0.05 - null hypothesis (Ho) rejected

-2.885087836
0.002640493

P(T<=t) two tail is

0.005280985

value < 0.05 - null hypothesis (Ho) rejectted

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists leaf No between Control and Camphor

* Sig dif in leaf no exists btween Cineole and Camphor

* Mean leaf No is reduced in Cineole

* Mean leaf No is reduced in Camphor

* Mean Cineole leaf no is reduced below Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-43

* Sig dif exists leaf No between Control and Cineole

Hymenosporum flavum - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

98.53846154

25.8

Variance

7572.102564

573.6666667

13

25

Observations
Pooled Variance

2906.478632

Hypothesized Mean difference

91.09302326

Mean

Variance

7572.102564

13537.46733

Variance

13

43

Observations

Hypothesized Mean difference

36

Cineole

98.53846154

Pooled Variance

df

Camphor

Mean

df

12211.83072

Observations
Pooled Variance

Hypothesized Mean difference

54

df

Camphor
25.8

91.09302326

573.6666667

13537.46733

25

43

8823.357999
0
66

t Stat

3.945757822

t Stat

0.212868682

t Stat

P(T<=t) one-tail

0.000176545

P(T<=t) one-tail

0.416115481

P(T<=t) one-tail

-2.763755397

t CritiCamphorl one-tail

1.688297289

t CritiCamphorl one-tail

1.673565748

t CritiCamphorl one-tail

1.668270215

0.003699028

P(T<=t) two-tail

0.00035309

P(T<=t) two-tail

0.832230963

P(T<=t) two-tail

0.007398056

t CritiCamphorl two-tail

2.02809133

t CritiCamphorl two-tail

2.004881026

t CritiCamphorl two-tail

1.996563697

P(T<=t) two tail is

0.00035309

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists between Control and Cineole


* Mean leaf area reduced in Cineole compared to Control

P(T<=t) two tail is

0.832230963

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* Not
exists
between
Control
andand
Camphor
*Nosig
sigdif
difininleaf
leafarea
area
exists
between
Control
Camphor

0.007398056

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif in leaf area exists between Cineoleand Camphor


* Mean leaf area is reduced in Cineole compared to Camphor

Lepiderema pulchella - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

32.23333333

33.08045977

Mean

32.23333333

35.07368421

Mean

33.08045977

35.07368421

Variance

26.04606742

22.19112537

Variance

26.04606742

25.47323628

Variance

22.19112537

25.47323628

90

87

90

95

87

95

Observations
Pooled Variance

24.15163875

Hypothesized Mean difference

t Stat

Pooled Variance

df

Observations

25.75182629

Hypothesized Mean difference

175
-1.146487122

t Stat

Pooled Variance

df

Observations

23.90511662

Hypothesized Mean difference

183

df

-3.805092066

t Stat

180
-2.747239393

P(T<=t) one-tail

0.126579419

P(T<=t) one-tail

9.65685E-05

P(T<=t) one-tail

0.003310618

t CritiCamphorl one-tail

1.653606887

t CritiCamphorl one-tail

1.653222625

t CritiCamphorl one-tail

1.653363597

P(T<=t) two-tail

0.253158838

P(T<=t) two-tail

0.000193137

P(T<=t) two-tail

0.006621236

t CritiCamphorl two-tail

1.973612598

t CritiCamphorl two-tail

1.973012331

t CritiCamphorl two-tail

P(T<=t) two tail is

0.253158838

P(T<=t) two tail is

0.000193137

value > 0.05 - null hypothesis (Ho) not rejected

1.97323061

P(T<=t) two tail is

0.006621236

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* No significant difference exists in shoot length between the Control and Cineole treatments.

* Their is a significant difference in shoot length between the Control and Camphor treatments.

* Their is a significant difference in shoot length between the Cineole and Camphor treatments.

* Mean length is similar in both treatments

* Mean length of the Camphor treatment is slightly higher than Control.

* Mean shoot length is reduced in the Cineole treatment.

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-44

t-Test: Two-Sample Assuming Equal Variances

Length
Lepiderema pulchella - Radical
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Mean

59.61111111

55.75862069

Mean

59.61111111

63.56842105

Variance

243.8358302

343.8131516

Variance

243.8358302

220.056439

90

87

90

95

Observations
Pooled Variance
Hypothesized Mean difference
df

292.9675424

Observations
Pooled Variance

Hypothesized Mean difference

175

df

231.6212795

55.75862069

63.56842105

Variance

343.8131516

220.056439

87

95

Observations
Pooled Variance

Hypothesized Mean difference

183

df

-1.767697839

Camphor

Mean

t Stat

279.1846461
0
180

t Stat

1.497014688

t Stat

P(T<=t) one-tail

0.068095429

P(T<=t) one-tail

0.039388953

P(T<=t) one-tail

0.000956775

t CritiCamphorl one-tail

1.653606887

t CritiCamphorl one-tail

1.653222625

t CritiCamphorl one-tail

1.653363597

P(T<=t) two-tail

0.136190858

P(T<=t) two-tail

0.078777906

P(T<=t) two-tail

0.00191355

t CritiCamphorl two-tail

1.973612598

t CritiCamphorl two-tail

1.973012331

t CritiCamphorl two-tail

1.97323061

P(T<=t) two tail is


P(T<=t) two tail is

0.136190858

P(T<=t) two tail is

-3.149777687

0.00191355

0.078777906
value < 0.05 - null hypothesis (Ho) rejected

value > 0.05 - null hypothesis (Ho) not rejected

value > 0.05 - null hypothesis (Ho) not rejected


* Radical length was significantly increased in the Camphor treatment compared to Cineole

*no significant effect on rad. length due to the Cineole treatment compared to Control.

*no significant effect on rad. length due to the Camphor treatment compared to Control.

Lepiderema pulchella - Leaf Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

1.944444444

1.876712329

Mean

1.944444444

1.909090909

Mean

1.876712329

1.909090909

Variance

0.109546166

0.192922374

Variance

0.109546166

0.162679426

Variance

0.192922374

0.162679426

72

73

72

77

73

77

Observations
Pooled Variance

0.151525795

Hypothesized Mean difference

Pooled Variance

df

Observations

0.137016423

Hypothesized Mean difference

143

Pooled Variance

df

Observations

0.177392212

Hypothesized Mean difference

147

df

148

t Stat

1.047598897

t Stat

0.582592667

t Stat

P(T<=t) one-tail

0.148295432

P(T<=t) one-tail

0.280530056

P(T<=t) one-tail

0.319309699

t CritiCamphorl one-tail

1.655284905

t CritiCamphorl one-tail

1.655214419

t CritiCamphorl one-tail

1.65558049

-0.470599749

P(T<=t) two-tail

0.296590865

P(T<=t) two-tail

0.561060113

P(T<=t) two-tail

0.638619399

t CritiCamphorl two-tail

1.976691237

t CritiCamphorl two-tail

1.976231943

t CritiCamphorl two-tail

1.976122803

P(T<=t) two tail is

0.296590865

P(T<=t) two tail is

0.561060113

value > 0.05 - null hypothesis (Ho) not rejected

P(T<=t) two tail is

0.638619399

value > 0.05 - null hypothesis (Ho) not rejected

value > 0.05 - null hypothesis (Ho) not rejected

* No significant difference in leaf No. exists between the Control and Camphor treatments.

* Their is no significant difference in Leaf No. between the Cineole and Camphor treatments.

* Mean length is similar in both treatments

* Mean length similar in Control and Camphor - no effect on leaf No. due Camphor treatment

* Mean Leaf No. is similar in both treatments.

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-45

* No significant difference in leaf No. exists between the Control and Cineole treatments.

Lepiderema pulchella - Leaf Area


t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

96.25555556

89.88311688

Mean

96.25555556

86.38947368

Mean

89.88311688

86.38947368

Variance

998.2598002

1185.236159

Variance

998.2598002

897.8573348

Variance

1185.236159

897.8573348

90

77

90

95

77

95

Observations
Pooled Variance
Hypothesized Mean difference
df

1084.382244

Observations
Pooled Variance

Hypothesized Mean difference

165

df

946.6869492

Pooled Variance

t Stat

1.246587729

t Stat

0.107157753

P(T<=t) one-tail

0.015270427

1026.332574

Hypothesized Mean difference

183

P(T<=t) one-tail

Observations

df

2.17991075

170

t Stat

0.711176561

P(T<=t) one-tail

0.238974741

t CritiCamphorl one-tail

1.654141215

t CritiCamphorl one-tail

1.653222625

t CritiCamphorl one-tail

1.653866093

P(T<=t) two-tail

0.214315506

P(T<=t) two-tail

0.030540854

P(T<=t) two-tail

0.477949483

t CritiCamphorl two-tail

1.974444785

t CritiCamphorl two-tail

1.973012331

t CritiCamphorl two-tail

1.974017323

P(T<=t) two tail is

0.214315506

value > 0.05 - null hypothesis (Ho) Not rejected

P(T<=t) two tail is

0.030540854

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.477949483

value > 0.05 - null hypothesis (Ho) not rejected

* No significant difference exists in leaf Area between the Control and Cineole treatments.

* A significant difference in leal Area exists between the Control and Camphor treatments.

* No significant difference in leaf Area exists between the Cineole and Camphor treatments.

* Mean length is similar for the Control and Cineole treatments

* Mean Area is significantly reduced in the Camphor treatment indicating effect.

* Mean Area is similar between the treatments.

*Ligustrum lucidum - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Cineole

Pooled Variance

Control

38.59375

26.04166667

Mean

117.7384868

214.8492908

Variance

96

48

Observations

149.8807952

Hypothesized Mean difference

Camphor
31.96875

117.7384868

169.5674342

96

96

143.6529605

Hypothesized Mean difference

142

Cineole

38.59375

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

26.04166667

31.96875

Variance

214.8492908

169.5674342

48

96

Observations
Pooled Variance

df

Camphor

Mean

184.555091

Hypothesized Mean difference

190

df

142

t Stat

5.799863983

t Stat

3.829562933

t Stat

P(T<=t) one-tail

2.06756E-08

P(T<=t) one-tail

8.71248E-05

P(T<=t) one-tail

0.007386545

t CritiCamphorl one-tail

1.655655524

t CritiCamphorl one-tail

1.652913397

t CritiCamphorl one-tail

1.655655524

P(T<=t) two-tail

4.13512E-08

P(T<=t) two-tail

P(T<=t) two-tail

0.014773091

t CritiCamphorl two-tail

1.976809472

t CritiCamphorl two-tail

t CritiCamphorl two-tail

1.976809472

P(T<=t) two tail is

4.13512E-08

0.00017425
1.972530299

P(T<=t) two tail is

0.00017425

value < 0.05 - null hypothesis (Ho) rejected

-2.468044623

P(T<=t) two tail is

0.014773091

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in shoot length between Control and Camphor

* Sig dif exists in shoot length between Cineole and Camphor

* Cineole Mean shoot length reduced below Control

* Cineole Mean shoot length reduced below Control

* Cineole Mean shoot length reduced below Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-46

* Sig dif exists in shoot length between Control and Cineole

*Ligustrum lucidum - Radical


Length
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Mean

48.47916667

32.08333333

Mean

48.47916667

38.8317757

Variance

142.2732456

207.2695035

Variance

142.2732456

134.7072827

96

48

96

107

Observations
Pooled Variance

163.7860915

Hypothesized Mean difference

Pooled Variance

df

Observations

138.2832353

Hypothesized Mean difference

142

38.8317757

Variance

207.2695035

134.7072827

48

107

Observations

156.9976381

Hypothesized Mean difference

201

Camphor

32.08333333

Pooled Variance

df

Cineole
Mean

df

153

t Stat

7.247195458

t Stat

5.835856434

t Stat

P(T<=t) one-tail

1.24803E-11

P(T<=t) one-tail

1.05849E-08

P(T<=t) one-tail

0.001150938

t CritiCamphorl one-tail

1.655655524

t CritiCamphorl one-tail

1.652470019

t CritiCamphorl one-tail

1.654873358

P(T<=t) two-tail

2.49606E-11

P(T<=t) two-tail

2.11697E-08

P(T<=t) two-tail

0.002301877

t CritiCamphorl two-tail

1.976809472

t CritiCamphorl two-tail

1.971834536

t CritiCamphorl two-tail

1.975590749

P(T<=t) two tail is

2.49606E-11

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

2.11697E-08

-3.100299443

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.002301877

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif in rad length exists between Control and Cineole

* Sig dif in rad length exists between Control and Camphor

* Sig dif in rad length exists between Cineole and Camphor

* Cineole mean rad length reduced below Control

* Camphor mean rad length reduced below Control

* Cineole mean rad length reduced below Camphor

*Ligustrum lucidum - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

2.458333333

1.208333333

Mean

2.458333333

1.855670103

Mean

1.208333333

1.855670103

Variance

1.219298246

1.147163121

Variance

1.219298246

1.187285223

Variance

1.147163121

1.187285223

96

48

96

97

48

97

Observations
Pooled Variance

1.195422535

Hypothesized Mean difference

Pooled Variance

df

Observations

1.203207931

Hypothesized Mean difference

142

Pooled Variance

df

Observations

1.174098239

Hypothesized Mean difference

191

df

143

t Stat

6.467318998

t Stat

3.816341072

t Stat

P(T<=t) one-tail

7.50449E-10

P(T<=t) one-tail

9.14345E-05

P(T<=t) one-tail

t CritiCamphorl one-tail

1.655655524

t CritiCamphorl one-tail

1.652870196

t CritiCamphorl one-tail

P(T<=t) two-tail

0.000182869

P(T<=t) two-tail

0.000917969

t CritiCamphorl two-tail

1.972462087

t CritiCamphorl two-tail

1.976691237

P(T<=t) two-tail
t CritiCamphorl two-tail

1.5009E-09
1.976809472

P(T<=t) two tail is

1.5009E-09

P(T<=t) two tail is

0.000182869

value < 0.05 - null hypothesis (Ho) rejected

-3.385326359
0.000458985
1.65558049

P(T<=t) two tail is

0.000917969

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists between Camphor and Control leaf No

* Sig dif exists between Cineole and Control leaf No

* Mean leadf no Cineole reduced below Control

* Mean leadf no Camphor reduced below Control

* Mean leadf no Cineole reduced below Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-47

* Sig dif exists between Cineole and Control leaf No

*Ligustrum lucidum - Leaf Area


t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

90.13541667

40.125

Variance

3121.402522

2633.81383

96

48

Observations
Pooled Variance

2960.017532

Hypothesized Mean difference

80.50515464

Mean

Variance

3121.402522

5403.273411

Variance

96

97

Observations

4268.31145

Hypothesized Mean difference

142

Cineole

90.13541667

Pooled Variance

df

Camphor

Mean

Pooled Variance

df

Observations

5.199820123

t Stat

1.023889012

t Stat

P(T<=t) one-tail

3.41151E-07

P(T<=t) one-tail

0.153591365

P(T<=t) one-tail

t CritiCamphorl one-tail

1.655655524

t CritiCamphorl one-tail

1.652870196

t CritiCamphorl one-tail

P(T<=t) two-tail

6.82302E-07

P(T<=t) two-tail

t CritiCamphorl two-tail

1.976809472

t CritiCamphorl two-tail

P(T<=t) two tail is

6.82302E-07

0.30718273
1.972462087

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in leaf area between Control and Cineole


* Cineole leaf area reduced below Control

0.30718273

5403.273411

48

97

df

t Stat

80.50515464

2633.81383

4493.031451

Hypothesized Mean difference

191

Camphor

40.125

143
-3.413663694
0.000417053
1.65558049

P(T<=t) two-tail

0.000834106

t CritiCamphorl two-tail

1.976691237

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists between Control and Camphor

0.000834106

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in leaf area between Cineole and Camphor


* Cineole leaf area reduced below Camphor

Lomandra longifolia - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

119.3064516

103.2386364

Mean

119.3064516

90.18518519

Mean

103.2386364

90.18518519

Variance

1220.642253

851.1722832

Variance

1220.642253

962.1527778

Variance

851.1722832

962.1527778

62

88

62

81

88

81

Observations
Pooled Variance

1003.453825

Hypothesized Mean difference

Pooled Variance

df

Observations

1073.981558

Hypothesized Mean difference

148

Pooled Variance

df

Observations

904.336592

Hypothesized Mean difference

141

df

167

t Stat

3.059142337

t Stat

5.266016084

t Stat

2.819037093

P(T<=t) one-tail

0.001317836

P(T<=t) one-tail

2.54735E-07

P(T<=t) one-tail

0.002699705

t CritiCamphorl one-tail

1.655214419

t CritiCamphorl one-tail

1.655732831

t CritiCamphorl one-tail

1.654029802

P(T<=t) two-tail

0.002635671

P(T<=t) two-tail

P(T<=t) two-tail

0.005399411

t CritiCamphorl two-tail

1.976122803

t CritiCamphorl two-tail

t CritiCamphorl two-tail

1.974271981

P(T<=t) two tail is

0.002635671

5.0947E-07
1.976932253

P(T<=t) two tail is

5.0947E-07

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.005399411

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Camphor

*Sig difference exists in shoot length between Cineole and Camphor

* Mean shoot length sig decreased in Camphor below Control

* Mean shoot length sig decreased in Camphor below Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-48

* Sig difference exists in shoot length between Control and Cineole


* Mean shoot length sig decreased in Cineole below Control

Lomandra longifolia - Radical


Length
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

66.69354839

61.875

Variance

325.3635643

423.1681034

62

88

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat
P(T<=t) one-tail

382.8567731
0

56.58536585

Mean

Variance

325.3635643

475.2333634

Variance

62

82

Observations

Hypothesized Mean difference


df

1.485215963
0.06980676

Cineole

66.69354839

Pooled Variance

148

Camphor

Mean

410.8526751

Observations
Pooled Variance

Hypothesized Mean difference

142

df

t Stat

2.963135702

t Stat

P(T<=t) one-tail

0.001785888

P(T<=t) one-tail

Camphor

61.875

56.58536585

423.1681034

475.2333634

88

82

448.2709967
0
168
1.627717815
0.05272936

t CritiCamphorl one-tail

1.655214419

t CritiCamphorl one-tail

1.655655524

t CritiCamphorl one-tail

1.653975232

P(T<=t) two-tail

0.139613519

P(T<=t) two-tail

0.003571775

P(T<=t) two-tail

0.105458719

t CritiCamphorl two-tail

1.976122803

t CritiCamphorl two-tail

1.976809472

t CritiCamphorl two-tail

1.974185579

P(T<=t) two tail is

0.139613519

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Cineole

0.003571775

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Camphor


* Mean rad length sig decreased in Camphor below Control

P(T<=t) two tail is

0.105458719

value > 0.05 - null hypothesis (Ho) accepted

*No Sig difference exists in rad. length between Cineole and Camphor

Lomandra longifolia - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

4.838709677

4.386363636

Mean

4.838709677

3.926829268

Mean

4.386363636

3.926829268

Variance

0.727657324

0.423719958

Variance

0.727657324

0.661246612

Variance

0.423719958

0.661246612

62

88

62

82

88

82

Observations
Pooled Variance

0.54899144

Hypothesized Mean difference

t Stat

Pooled Variance

df

0.689775158

Hypothesized Mean difference

148

0.00016185

Observations
Pooled Variance

df

3.681964175

P(T<=t) one-tail

Observations

0.538241738

Hypothesized Mean difference

142

df

168

t Stat

6.523875068

t Stat

P(T<=t) one-tail

5.61782E-10

P(T<=t) one-tail

4.080873315
3.46094E-05

1.655655524

t CritiCamphorl one-tail

1.653975232

t CritiCamphorl one-tail

1.655214419

t CritiCamphorl one-tail

P(T<=t) two-tail

0.000323701

P(T<=t) two-tail

1.12356E-09

P(T<=t) two-tail

6.92188E-05

t CritiCamphorl two-tail

1.976122803

t CritiCamphorl two-tail

1.976809472

t CritiCamphorl two-tail

1.974185579

P(T<=t) two tail is

0.000323701

P(T<=t) two tail is

1.12356E-09

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

6.92188E-05

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf No. between Control and Camphor

* Sig difference exists in leaf No. between Cineole and Camphor

* Mean leaf no. sig reduced in Camphor below Control

* Mean leaf no. sig reduced in Camphor below Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-49

* Sig difference exists in leaf No. between Control and Cineole


* Mean leaf no. sig reduced in Cineole below Control

Lomandra longifolia - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

120.2741935

103.0113636

Mean

120.2741935

89.17073171

Mean

103.0113636

89.17073171

Variance

1279.415389

861.8044671

Variance

1279.415389

1068.365553

Variance

861.8044671

1068.365553

62

88

62

82

88

82

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat
P(T<=t) one-tail
t CritiCamphorl one-tail
P(T<=t) two-tail
t CritiCamphorl two-tail

1033.927887

Observations
Pooled Variance

Hypothesized Mean difference

148

df

3.237862621
0.00074331
1.655214419
0.00148662
1.976122803

P(T<=t) two tail is

0.00148662

value < 0.05 - null hypothesis (Ho) rejected

1159.027806

Observations
Pooled Variance

Hypothesized Mean difference

142

df

961.396419
0
168

t Stat

5.428543551

t Stat

2.90822813

P(T<=t) one-tail

1.19728E-07

P(T<=t) one-tail

0.00206318

t CritiCamphorl one-tail

1.655655524

t CritiCamphorl one-tail

1.653975232

P(T<=t) two-tail

2.39456E-07

P(T<=t) two-tail

0.004126361

t CritiCamphorl two-tail

1.976809472

t CritiCamphorl two-tail

1.974185579

P(T<=t) two tail is

2.39456E-07

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.004126361

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Control and Cineole

* Sig difference exists in leaf area between Control and Camphor

* Sig difference exists in leaf area between Cineole and Camphor

* Mean leaf area sig reduced in Cineole compared to Control

* Mean leaf area sig reduced in Camphor compared to Control

* Mean leaf area sig reduced in Camphor compared to Cineole

Lophostemon confertus - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

13.54285714

10.10714286

Mean

13.54285714

8.4

Variance

9.137815126

5.876984127

Variance

9.137815126

8.685714286

35

28

35

15

Observations
Pooled Variance

7.694496487

Hypothesized Mean difference

Pooled Variance

df

Observations

9.005952381

Hypothesized Mean difference

61

10.10714286

8.4

Variance

5.876984127

8.685714286

28

15

Observations
Pooled Variance

df

6.836062718

Hypothesized Mean difference

48

Camphor

Mean

df

41

t Stat

4.885057736

t Stat

5.553084565

t Stat

P(T<=t) one-tail

3.91134E-06

P(T<=t) one-tail

5.99133E-07

P(T<=t) one-tail

t CritiCamphorl one-tail

1.670218808

t CritiCamphorl one-tail

1.677224191

t CritiCamphorl one-tail

1.682878974

P(T<=t) two-tail

7.82269E-06

P(T<=t) two-tail

1.19827E-06

P(T<=t) two-tail

0.047765579

t CritiCamphorl two-tail

1.999624146

t CritiCamphorl two-tail

P(T<=t) two tail is

7.82269E-06

2.01063358

P(T<=t) two tail is

t CritiCamphorl two-tail

1.19827E-06

value < 0.05 - null hypothesis (Ho) rejected

2.040597092
0.02388279

2.01954208

P(T<=t) two tail is

0.047765579

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* sig dif in shoot length exists between Control and Camphor

* sig dif in shoot length exists between Cineole and Camphor

* Mean shoot length reduced in Cineole

* Mean shoot length reduced in Camphor

* Mean shoot length reduced in Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-50

* sig dif in shoot length exists between Control and Cineole

Radicle
Length
Lophostemon confertus - Radical
Length
t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Cineole
18.39285714

Mean

85.71764706

62.61772487

Variance

35

28

Observations
Pooled Variance

Control

26.4

75.49309133

Hypothesized Mean difference

Pooled Variance

df

Observations

Camphor

df

Camphor

15.93333333

Mean

18.39285714

15.93333333

85.71764706

136.7809524

Variance

62.61772487

136.7809524

35

15

28

15

100.6111111

Hypothesized Mean difference

61

Cineole

26.4

Observations
Pooled Variance

Hypothesized Mean difference

48

df

87.94175377
0
41

t Stat

3.634684637

t Stat

3.381271705

t Stat

0.819679149

P(T<=t) one-tail

0.000286412

P(T<=t) one-tail

0.000721205

P(T<=t) one-tail

0.208569364

t CritiCamphorl one-tail

1.670218808

t CritiCamphorl one-tail

1.677224191

t CritiCamphorl one-tail

1.682878974

P(T<=t) two-tail

0.000572823

P(T<=t) two-tail

0.00144241

P(T<=t) two-tail

0.417138729

t CritiCamphorl two-tail

1.999624146

t CritiCamphorl two-tail

2.01063358

t CritiCamphorl two-tail

P(T<=t) two tail is

0.000572823

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.00144241

value < 0.05 - null hypothesis (Ho) rejected

* sig dif in rad length exists between Control and Cineole

* sig dif in rad length exists between Control and Camphor

* Mean rad length reduced in Cineole

* Mean rad length reduced in Camphor

2.01954208

P(T<=t) two tail is

0.417138729

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif in rad length exists between Cineole and Camphor

Lophostemon confertus - Leaf Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

1.885714286

1.357142857

Mean

1.885714286

1.2

Variance

0.221848739

0.904761905

Variance

0.221848739

1.028571429

35

28

35

15

Observations
Pooled Variance

0.52412178

Hypothesized Mean difference

Pooled Variance

df

Observations

0.457142857

Hypothesized Mean difference

61

1.357142857

1.2

Variance

0.904761905

1.028571429

28

15

Observations
Pooled Variance

df

0.947038328

Hypothesized Mean difference

48

Camphor

Mean

df

41

t Stat

2.879587924

t Stat

3.286335345

t Stat

P(T<=t) one-tail

0.002742707

P(T<=t) one-tail

0.000950922

P(T<=t) one-tail

t CritiCamphorl one-tail

1.670218808

t CritiCamphorl one-tail

1.677224191

t CritiCamphorl one-tail

1.682878974

P(T<=t) two-tail

0.005485413

P(T<=t) two-tail

0.001901844

P(T<=t) two-tail

0.616497461

t CritiCamphorl two-tail

1.999624146

t CritiCamphorl two-tail

P(T<=t) two tail is

0.005485413

2.01063358

P(T<=t) two tail is

t CritiCamphorl two-tail

0.001901844

value < 0.05 - null hypothesis (Ho) rejected

0.50466279
0.30824873

2.01954208

P(T<=t) two tail is

0.616497461

value < 0.05 - null hypothesis (Ho) rejected

* sig dif in leaf no exists between Control and Camphor

* Mean leaf No reduced in Cineole

* Mean rad leaf No reduced in Camphor

* No sig dif in leaf No exists between Cineole and Camphor

A11-51

* sig dif in Leaf No exists between Control and Cineole

value > 0.05 - null hypothesis (Ho) accepted

Lophostemon confertus - Leaf Area


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

1.971428571

0.678571429

Mean

1.971428571

0.6

Variance

2.322689076

0.226190476

Variance

2.322689076

0.257142857

35

28

35

15

Observations
Pooled Variance

1.394730679

Hypothesized Mean difference

Pooled Variance

df

Observations

1.720238095

Hypothesized Mean difference

61

0.678571429

0.6

Variance

0.226190476

0.257142857

28

15

Observations
Pooled Variance

df

Hypothesized Mean difference

48

Camphor

Mean

df

0.236759582
0
41

t Stat

4.317656869

t Stat

3.388235294

t Stat

P(T<=t) one-tail

2.94603E-05

P(T<=t) one-tail

0.000706616

P(T<=t) one-tail

t CritiCamphorl one-tail

1.670218808

t CritiCamphorl one-tail

1.677224191

t CritiCamphorl one-tail

1.682878974

P(T<=t) two-tail

5.89206E-05

P(T<=t) two-tail

0.001413233

P(T<=t) two-tail

0.616497461

t CritiCamphorl two-tail

1.999624146

t CritiCamphorl two-tail

P(T<=t) two tail is

5.89206E-05

2.01063358

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

t CritiCamphorl two-tail

0.001413233

value < 0.05 - null hypothesis (Ho) rejected

* sig dif in leaf area exists between Control and Cineole

* sig dif in leaf area exists between Control and Camphor

* Mean leaf area reduced in Cineole

* Mean leaf area reduced in Camphor

0.50466279
0.30824873

2.01954208

P(T<=t) two tail is

0.616497461

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif in leaf No exists between Cineole and Camphor

Macaranga tanarius - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

80.56818182

68.02777778

Mean

80.56818182

78.26984127

Mean

68.02777778

78.26984127

Variance

426.7161734

102.3372457

Variance

426.7161734

283.5873016

Variance

102.3372457

283.5873016

44

72

44

63

72

63

Observations
Pooled Variance

224.6907009

Hypothesized Mean difference

Pooled Variance

df

Observations

342.2019824

Hypothesized Mean difference

114

Pooled Variance

df

Observations

186.829753

Hypothesized Mean difference

105

df

133

t Stat

4.372024346

t Stat

0.632379587

t Stat

P(T<=t) one-tail

1.36651E-05

P(T<=t) one-tail

0.264256763

P(T<=t) one-tail

-4.343442561
1.37991E-05

t CritiCamphorl one-tail

1.658329438

t CritiCamphorl one-tail

1.659495865

t CritiCamphorl one-tail

1.656389941

P(T<=t) two-tail

2.73302E-05

P(T<=t) two-tail

0.528513526

P(T<=t) two-tail

2.75983E-05

t CritiCamphorl two-tail

1.980993147

t CritiCamphorl two-tail

1.982816684

t CritiCamphorl two-tail

1.977959982

P(T<=t) two tail is

2.73302E-05

P(T<=t) two tail is

0.528513526

value < 0.05 - null hypothesis (Ho) rejected

*Sig. dif exists i shoot length between Control andCineole

P(T<=t) two tail is

2.75983E-05

value > 0.05 - null hypothesis (Ho) accepted

* No Sig.dif exist in shoot length between Control and Camphor

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in shoot length between Cineole and Camphor


* Cineole is reduced below Camphor

A11-52

* Mean shoot length reduced in Cineole below Control

Radicle
Length
Length
Macaranga tanarius - Radical
t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

42.52272727

42.88888889

Mean

42.52272727

43.85714286

Mean

42.88888889

43.85714286

Variance

32.81342495

141.5931142

Variance

32.81342495

403.7373272

Variance

141.5931142

403.7373272

44

72

44

63

72

63

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

100.5621788

Observations
Pooled Variance

251.8351577

Hypothesized Mean difference

114

df

-0.190817882

t Stat

Observations
Pooled Variance

263.7956797

Hypothesized Mean difference

105

df

-0.427993969

t Stat

0
133
-0.345561289

P(T<=t) one-tail

0.424503845

P(T<=t) one-tail

0.334766048

P(T<=t) one-tail

0.365109202

t CritiCamphorl one-tail

1.658329438

t CritiCamphorl one-tail

1.659495865

t CritiCamphorl one-tail

1.656389941

P(T<=t) two-tail

0.849007689

P(T<=t) two-tail

0.669532097

P(T<=t) two-tail

0.730218403

t CritiCamphorl two-tail

1.980993147

t CritiCamphorl two-tail

1.982816684

t CritiCamphorl two-tail

1.977959982

P(T<=t) two tail is

0.849007689

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

*No Sig.dif exists in rad.length between Control and Cineole

0.669532097

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig. dif in rad length between Control and Camphor

0.730218403

value > 0.05 - null hypothesis (Ho) accepted

*No sig dif in rad length between Cineole and Camphor

Macaranga tanarius - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

0.931818182

1.083333333

Mean

0.931818182 1.063492063

Mean

1.083333333

1.063492063

Variance

0.111522199

0.133802817

Variance

0.111522199 0.350742448

Variance

0.133802817

0.350742448

44

72

72

63

Observations
Pooled Variance

0.125398724

Hypothesized Mean difference

44

Pooled Variance

df

Observations

Hypothesized Mean difference

114

63

0.25277606

Pooled Variance

df

Observations

0.23493257

Hypothesized Mean difference

105

df

133

t Stat

-2.236006684

t Stat

-1.333019163

t Stat

0.237283472

P(T<=t) one-tail

0.013649068

P(T<=t) one-tail

0.092705331

P(T<=t) one-tail

0.406400999

t CritiCamphorl one-tail

1.658329438

t CritiCamphorl one-tail

1.659495865

t CritiCamphorl one-tail

1.656389941

P(T<=t) two-tail

0.027298137

P(T<=t) two-tail

0.185410662

P(T<=t) two-tail

0.812801998

t CritiCamphorl two-tail

1.980993147

t CritiCamphorl two-tail

1.982816684

t CritiCamphorl two-tail

1.977959982

P(T<=t) two tail is

0.027298137

P(T<=t) two tail is

0.185410662

value < 0.05 - null hypothesis (Ho) rejected

0.812801998

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

*No sig dif in leaf number exists between Control and Camphor

*No sig dif in leaf number exists between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-53

* Sig.dif exists in leaf No.between Control and Cineole

P(T<=t) two tail is

* Cineole leaf No. is increased against Control

Macaranga tanarius - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Cineole
72.92957746

3694.256757

17338.0664

38

71

Observations
Pooled Variance

Control

27.5

12620.11353

Hypothesized Mean difference

Variance

Cineole

3694.256757 15349.03738
38

63

10993.21028

Hypothesized Mean difference

107

Camphor
27.5 66.79365079

Observations
Pooled Variance

df

Mean

Variance

-2.011938912

t Stat

-1.824574075

t Stat

P(T<=t) one-tail

0.023369724

P(T<=t) one-tail

0.035540656

P(T<=t) one-tail

t CritiCamphorl one-tail

1.659218469

t CritiCamphorl one-tail

1.660391717

t CritiCamphorl one-tail

P(T<=t) two-tail

0.046739448

P(T<=t) two-tail

0.071081312

P(T<=t) two-tail

t CritiCamphorl two-tail

1.982384674

t CritiCamphorl two-tail

1.984217306

t CritiCamphorl two-tail

0.046739448

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif in leaf area exists between Control and Cineole


* Cineole is increased above Control

15349.03738

71

63

16403.8255
0

df

t Stat

P(T<=t) two tail is

66.79365079

17338.0664

Hypothesized Mean difference

99

0.071081312

132
0.276792705
0.39118612
1.656478616
0.78237224
1.978096407

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif in leaf No between Control and Camphor

Camphor

72.92957746

Observations
Pooled Variance

df

Mean

0.78237224

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif in leaf No between Cineole and Camphor

Omalanthus populifolius - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Cineole

Pooled Variance

Control

77.6835443

70.90789474

Mean

363.1165206

193.4714035

Variance

79

76

Observations

279.9571495

Hypothesized Mean difference

Cineole

Camphor

68.71794872

Mean

70.90789474

68.71794872

363.1165206

229.5038295

Variance

193.4714035

229.5038295

79

78

76

78

296.7411837

Hypothesized Mean difference

153

Camphor

77.6835443

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

Pooled Variance

df

Observations

211.7246719

Hypothesized Mean difference

155

df

152

t Stat

2.520342992

t Stat

3.260627999

t Stat

0.933773216

P(T<=t) one-tail

0.006374644

P(T<=t) one-tail

0.000683423

P(T<=t) one-tail

0.175950934

t CritiCamphorl one-tail

1.654873358

t CritiCamphorl one-tail

1.654743755

t CritiCamphorl one-tail

1.654939297

P(T<=t) two-tail

0.012749288

P(T<=t) two-tail

0.001366847

P(T<=t) two-tail

0.351901867

t CritiCamphorl two-tail

1.975590749

t CritiCamphorl two-tail

1.975386112

t CritiCamphorl two-tail

1.975695341

P(T<=t) two tail is

0.012749288

P(T<=t) two tail is

0.001366847

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.351901867

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in shoot length between Control and Camphor

* Mean shoot length reduced by Cineole

* Mean shoot length reduced by Camphor

* No sig dif between Cineole and Camphor

A11-54

* Sig dif exists in shoot length between Control and Cineole

value > 0.05 - null hypothesis (Ho) accepted

Radicle
Length
Omalanthus populifolius - Radical
Length
t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Cineole

Pooled Variance

41.13333333

Mean

101.881001

110.5225225

Variance

89

75

105.8283627

Hypothesized Mean difference


df
t Stat

Control

37.73033708

Observations

t-Test: Two-Sample Assuming Equal Variances

41.13333333

37.17948718

101.881001

84.14918415

Variance

110.5225225

84.14918415

89

78

75

78

93.60615315

t Stat

Observations
Pooled Variance

df

-2.110396916

Camphor

Mean

Hypothesized Mean difference

162

Cineole

37.17948718

Observations
Pooled Variance

Camphor

37.73033708

97.07386653

Hypothesized Mean difference

165

df

0.367083737

t Stat

2.481423096

P(T<=t) one-tail

0.007091007

P(T<=t) one-tail

0.018179972

P(T<=t) one-tail

t CritiCamphorl one-tail

1.654314019

t CritiCamphorl one-tail

1.654141215

t CritiCamphorl one-tail

1.655007509

P(T<=t) two-tail

0.036359943

P(T<=t) two-tail

0.714026919

P(T<=t) two-tail

0.014182013

t CritiCamphorl two-tail

1.974717634

t CritiCamphorl two-tail

1.974444785

t CritiCamphorl two-tail

1.975799933

P(T<=t) two tail is

0.036359943

0.35701346

151

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif in rad length exists between Control and Cineole


* Mean rad length increased by Cineole compared to Control

0.714026919

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif in rad length between Control and Camphor

0.014182013

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif in rad length exists between Cineole and Camphor


* Camphor rad length reduced below Cineole

Omalanthus populifolius - Leaf Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

1.897435897

1.733333333

Mean

1.897435897

1.794871795

Mean

1.733333333

1.794871795

Variance

0.197136197

0.468468468

Variance

0.197136197

0.372960373

Variance

0.468468468

0.372960373

78

75

78

78

75

78

Observations
Pooled Variance

0.330106979

Hypothesized Mean difference

t Stat

Pooled Variance

df

0.285048285

Hypothesized Mean difference

151

0.03969825

Observations
Pooled Variance

df

1.766120093

P(T<=t) one-tail

Observations

0.419765665

Hypothesized Mean difference

154

df

151

t Stat

1.199688433

t Stat

-0.58732082

P(T<=t) one-tail

0.116051407

P(T<=t) one-tail

0.278932724

t CritiCamphorl one-tail

1.655007509

t CritiCamphorl one-tail

t CritiCamphorl one-tail

1.655007509

P(T<=t) two-tail

0.079396499

P(T<=t) two-tail

0.232102814

P(T<=t) two-tail

0.557865449

t CritiCamphorl two-tail

1.975799933

t CritiCamphorl two-tail

1.975486157

t CritiCamphorl two-tail

1.975799933

P(T<=t) two tail is

0.079396499

1.65480742

P(T<=t) two tail is

0.232102814

value > 0.05 - null hypothesis (Ho) accepted

0.557865449

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif in leaf no exists for Control and Camphor

* No sig dif in leaf no exists for Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-55

* No sig dif in leaf no exists for Control and Cineole

P(T<=t) two tail is

Omalanthus populifolius - Leaf Area


t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

117.7721519

95.13333333

Mean

117.7721519

79.98717949

Mean

95.13333333

79.98717949

Variance

5560.383317

5183.117117

Variance

5560.383317

3784.376457

Variance

5183.117117

3784.376457

79

75

79

78

75

78

Observations
Pooled Variance
Hypothesized Mean difference
df

5376.714246

Observations
Pooled Variance

4678.108941

Hypothesized Mean difference

152

df

Observations
Pooled Variance

Hypothesized Mean difference

155

df

4469.852012
0
151

t Stat

1.915045504

t Stat

3.460947127

t Stat

1.400838306

P(T<=t) one-tail

0.028682419

P(T<=t) one-tail

0.000347804

P(T<=t) one-tail

0.081657068

t CritiCamphorl one-tail

1.654939297

t CritiCamphorl one-tail

1.654743755

t CritiCamphorl one-tail

1.655007509

P(T<=t) two-tail

0.057364837

P(T<=t) two-tail

0.000695609

P(T<=t) two-tail

0.163314135

t CritiCamphorl two-tail

1.975695341

t CritiCamphorl two-tail

1.975386112

t CritiCamphorl two-tail

1.975799933

P(T<=t) two tail is

0.057364837

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in leaf area between Control and Cineole

0.000695609

value < 0.05 - null hypothesis (Ho) rejected

*Sig dif in leaf area exists between Control and Camphor


* Mean leaf area is reduced in Camphor

P(T<=t) two tail is

0.163314135

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in leaf area between Cineole and Camphor

Oplismenus aemulus - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean

Cineole

Control

28.93506494

23.38043478

28.3773069

34.3262064

77

92

Variance
Observations
Pooled Variance

31.6189228

Hypothesized Mean difference

Mean

Cineole

24.41836735

Mean

28.3773069

24.67883442

Variance

77

98

Variance
Observations

26.3035969

Hypothesized Mean difference

167

Camphor

28.93506494

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

df

24.41836735

34.3262064

24.67883442

92

98

Observations
Pooled Variance

29.34857299

Hypothesized Mean difference

173

Camphor

23.38043478

df

188

t Stat

6.395544663

t Stat

5.782995037

t Stat

-1.31979292

P(T<=t) one-tail

7.71753E-10

P(T<=t) one-tail

1.67982E-08

P(T<=t) one-tail

0.094254504

t CritiCamphorl one-tail

1.654029802

t CritiCamphorl one-tail

1.653709205

t CritiCamphorl one-tail

1.652999799

P(T<=t) two-tail

1.54351E-09

P(T<=t) two-tail

3.35963E-08

P(T<=t) two-tail

0.188509009

t CritiCamphorl two-tail

1.974271981

t CritiCamphorl two-tail

1.973771759

t CritiCamphorl two-tail

1.972662176

P(T<=t) two tail is

1.54351E-09

P(T<=t) two tail is

3.35963E-08

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.188509009

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Camphor


* Mean shoot length sig decreased in Camphor below Control

*No Sig difference exists in shoot length between Cineole and Camphor

A11-56

* Sig difference exists in shoot length between Control and Cineole


* Mean shoot length sig decreased in Cineole below Control

value > 0.05 - null hypothesis (Ho) accepted

Radicle
Length
Oplismenus aemulus - Radical
Length
t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

57.66233766

54.07608696

Mean

57.66233766

68.06122449

Variance

472.0950103

502.1589823

Variance

472.0950103

1485.89312

77

92

77

98

Observations
Pooled Variance

488.4771747

Hypothesized Mean difference


df

Observations
Pooled Variance

Hypothesized Mean difference

167

df

1040.525164

54.07608696

68.06122449

Variance

502.1589823

1485.89312

92

98

Observations
Pooled Variance

Hypothesized Mean difference

173

df

0
188

t Stat

1.050544451

t Stat

0.147492852

P(T<=t) one-tail

0.017849137

P(T<=t) one-tail

0.001387173

t CritiCamphorl one-tail

1.654029802

t CritiCamphorl one-tail

1.653709205

t CritiCamphorl one-tail

1.652999799

P(T<=t) two-tail

0.294985704

P(T<=t) two-tail

0.035698274

P(T<=t) two-tail

0.002774346

t CritiCamphorl two-tail

1.974271981

t CritiCamphorl two-tail

1.973771759

t CritiCamphorl two-tail

1.972662176

0.294985704

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

t Stat

1009.723936

P(T<=t) one-tail

P(T<=t) two tail is

-2.116898007

Camphor

Mean

0.035698274

value < 0.05 - null hypothesis (Ho) rejected

-3.031765223

P(T<=t) two tail is

0.002774346

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Cineole

* Sig difference exists in rad. length between Control and Camphor

*Sig difference exists in rad. length between Cineole and Camphor

* Mean rad length sig decreased in Cineole below Control

* Mean rad length sig increased in Camphor above Control

* Mean rad length sig increased in Camphor above Cineole

Oplismenus aemulus - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

2.415584416

2.065217391

Mean

2.415584416

2.255102041

Mean

2.065217391

2.255102041

Variance

0.377648667

0.281414238

Variance

0.377648667

0.212602567

Variance

0.281414238

0.212602567

77

92

77

98

92

98

Observations
Pooled Variance

0.325209547

Hypothesized Mean difference

df

Observations
Pooled Variance

0.285108368

Hypothesized Mean difference

167

Pooled Variance

df

Observations

0.245910344

Hypothesized Mean difference

173

df

188

t Stat

3.977750192

t Stat

1.973611814

t Stat

-2.63773772

P(T<=t) one-tail

5.17351E-05

P(T<=t) one-tail

0.025009155

P(T<=t) one-tail

0.004522518

t CritiCamphorl one-tail

1.654029802

t CritiCamphorl one-tail

1.653709205

t CritiCamphorl one-tail

1.652999799

P(T<=t) two-tail
t CritiCamphorl two-tail

0.00010347

P(T<=t) two-tail

1.974271981

t CritiCamphorl two-tail

P(T<=t) two tail is

0.00010347

0.05001831
1.973771759

P(T<=t) two tail is

0.05001831

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf No. between Control and Cineole

P(T<=t) two-tail

0.009045035

t CritiCamphorl two-tail

1.972662176

P(T<=t) two tail is

0.009045035

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf No. between Control and Camphor

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf No. between Cineole and Camphor


* Mean leaf no.reduced in Cineole Controlm[ared to Camphor

A11-57

* Mean leaf no. sig reduced in Cineole compared to Control

Oplismenus aemulus - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean difference
df

Cineole

Control

19.79220779

15.95121951

96.6404648

34.0469738

77

82

64.34700766

Mean
Variance
Observations
Pooled Variance

Hypothesized Mean difference

157

df

t Stat

3.017397532

t Stat

P(T<=t) one-tail

0.001487724

P(T<=t) one-tail

t CritiCamphorl one-tail

1.654616426

t CritiCamphorl one-tail

P(T<=t) two-tail

0.002975447

t CritiCamphorl two-tail

1.975190571

P(T<=t) two tail is

t-Test: Two-Sample Assuming Equal Variances

0.002975447

value < 0.05 - null hypothesis (Ho) rejected

Camphor

Cineole

19.79220779

14.31632653

Mean

96.6404648

36.96076162

Variance

77

98

63.17843469

Observations
Pooled Variance

Hypothesized Mean difference

173

df

4.523858034

t Stat

5.6228E-06

Camphor

15.95121951

14.31632653

34.0469738

36.96076162

82

98

35.63482447
0
178
1.829935275

P(T<=t) one-tail

0.034466164

1.653709205

t CritiCamphorl one-tail

1.653459094

P(T<=t) two-tail

1.12456E-05

P(T<=t) two-tail

0.068932327

t CritiCamphorl two-tail

1.973771759

t CritiCamphorl two-tail

1.973380677

P(T<=t) two tail is

1.12456E-05

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Control and Cineole

* Sig difference exists in leaf area between Control and Camphor

* Mean leaf area sig reduced in Cineole compared to Control

* Mean leaf area sig reduced in Camphor compared to Control

P(T<=t) two tail is

0.068932327

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Cineole and Camphor

*Paspalum dilatatum - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Control

34.5

31.78947368

Mean

41.11764706

41.09774436

Variance

52

57

Observations
Pooled Variance

Cineole

41.10723069

Hypothesized Mean difference

Cineole

Camphor

28.71428571

Mean

31.78947368

28.71428571

41.11764706

13.57142857

Variance

41.09774436

13.57142857

52

57

38.21804511

Hypothesized Mean difference

107

Camphor
34.5

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

Pooled Variance

df

Observations

38.43390735

Hypothesized Mean difference

57

df

62

t Stat

2.204552109

t Stat

2.324595725

t Stat

1.238542342

P(T<=t) one-tail

0.014813269

P(T<=t) one-tail

0.011839619

P(T<=t) one-tail

0.110091975

t CritiCamphorl one-tail

1.659218469

t CritiCamphorl one-tail

1.672028702

t CritiCamphorl one-tail

1.669804988

P(T<=t) two-tail

0.029626538

P(T<=t) two-tail

0.023679238

P(T<=t) two-tail

0.220183949

t CritiCamphorl two-tail

1.982384674

t CritiCamphorl two-tail

2.002466317

t CritiCamphorl two-tail

P(T<=t) two tail is

0.029626538

P(T<=t) two tail is

0.023679238

value < 0.05 - null hypothesis (Ho) rejected

1.99896931

P(T<=t) two tail is

0.220183949

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Camphor


* Mean shoot length sig decreased in Camphor below Control

*No Sig difference exists in shoot length between Cineole and Camphor

A11-58

* Sig difference exists in shoot length between Control and Cineole


* Mean shoot length sig decreased in Cineole below Control

value > 0.05 - null hypothesis (Ho) accepted

Length
*Paspalum dilatatum - Radical
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

39.80769231

40.70175439

Variance

126.4328808

169.141604

52

57

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

t-Test: Two-Sample Assuming Equal Variances

148.7851098
0

39.25

Variance

126.4328808

591.6428571

52

Observations

Hypothesized Mean difference


df

182.5789125

Mean
Variance
Observations
Pooled Variance

Hypothesized Mean difference

58

df

Camphor

40.70175439

39.25

169.141604

591.6428571

57

216.0861877
0
63

t Stat

0.108677687

t Stat

P(T<=t) one-tail

0.351528022

P(T<=t) one-tail

0.456916627

P(T<=t) one-tail

t CritiCamphorl one-tail

1.659218469

t CritiCamphorl one-tail

1.671553491

t CritiCamphorl one-tail

1.669402536

P(T<=t) two-tail

0.703056044

P(T<=t) two-tail

0.913833254

P(T<=t) two-tail

0.794498179

t CritiCamphorl two-tail

1.982384674

t CritiCamphorl two-tail

2.001715984

t CritiCamphorl two-tail

1.998341759

P(T<=t) two tail is

-0.382220336

Cineole

39.80769231

Pooled Variance

107

Camphor

Mean

0.703056044

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Cineole

P(T<=t) two tail is

0.913833254

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Camphor

P(T<=t) two tail is

0.261580512
0.39724909

0.794498179

value > 0.05 - null hypothesis (Ho) accepted

*No Sig difference exists in rad. length between Cineole and Camphor

*Paspalum dilatatum - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

2.269230769

1.842105263

Mean

2.269230769

1.142857143

Mean

1.842105263

1.142857143

Variance

0.279034691

0.171052632

Variance

0.279034691

0.142857143

Variance

0.171052632

0.142857143

52

57

52

57

Observations
Pooled Variance

0.222520716

Hypothesized Mean difference

t Stat

Pooled Variance

df

Observations

0.264700212

Hypothesized Mean difference

107
4.721675459

Pooled Variance

df

Observations

0.168324036

Hypothesized Mean difference

57

df

62

t Stat

5.437884944

t Stat

P(T<=t) one-tail

5.86794E-07

P(T<=t) one-tail

3.58899E-05

1.659218469

t CritiCamphorl one-tail

1.672028702

t CritiCamphorl one-tail

1.669804988

P(T<=t) two-tail

7.12981E-06

P(T<=t) two-tail

1.17359E-06

P(T<=t) two-tail

7.17799E-05

t CritiCamphorl two-tail

1.982384674

t CritiCamphorl two-tail

2.002466317

t CritiCamphorl two-tail

P(T<=t) one-tail
t CritiCamphorl one-tail

3.5649E-06

P(T<=t) two tail is

7.12981E-06

P(T<=t) two tail is

1.17359E-06

value < 0.05 - null hypothesis (Ho) rejected

4.255540231

1.99896931

P(T<=t) two tail is

7.17799E-05

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf No. between Control and Camphor

* Sig difference exists in leaf No. between Cineole and Camphor

* Mean leaf no. sig reduced in Cineole below Control

* Mean leaf no. sig reduced in Camphor below Control

* Mean leaf no. sig reduced in Camphor below Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-59

* Sig difference exists in leaf No. between Control and Cineole

*Paspalum dilatatum - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

41.30769231

38.45614035

Mean

41.30769231

49.57142857

Mean

38.45614035

49.57142857

Variance

337.8642534

301.0382206

Variance

337.8642534

272.6190476

Variance

301.0382206

272.6190476

52

57

52

57

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

318.5908156

Observations
Pooled Variance

Hypothesized Mean difference

107

df

0.8330873

330.996337

Observations
Pooled Variance

Hypothesized Mean difference

57

df

298.287978
0
62

t Stat

-1.12820895

t Stat

P(T<=t) one-tail

0.203325614

P(T<=t) one-tail

0.131979058

P(T<=t) one-tail

0.056574623

t CritiCamphorl one-tail

1.659218469

t CritiCamphorl one-tail

1.672028702

t CritiCamphorl one-tail

1.669804988

P(T<=t) two-tail

0.406651228

P(T<=t) two-tail

0.263958116

P(T<=t) two-tail

0.113149245

t CritiCamphorl two-tail

1.982384674

t CritiCamphorl two-tail

2.002466317

t CritiCamphorl two-tail

P(T<=t) two tail is

0.406651228

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Cineole

P(T<=t) two tail is

0.263958116

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Camphor

P(T<=t) two tail is

-1.606938716

1.99896931

0.113149245

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Cineole and Camphor

*Pennisetum clandestinum - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Control

65.75

41.11494253

Mean

136.8762626

101.0098904

Variance

100

87

Observations
Pooled Variance

Cineole

120.2032463

Hypothesized Mean difference

t Stat

Camphor

Mean

41.11494253

44.44578313

136.8762626

58.05495151

Variance

101.0098904

58.05495151

100

83

87

83

101.1671604

Observations
Pooled Variance

df

15.3261935

Cineole

44.44578313

Hypothesized Mean difference

185

Camphor
65.75

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

80.04378928

Hypothesized Mean difference

181

df

168

t Stat

14.26459007

t Stat

P(T<=t) one-tail

4.61528E-35

P(T<=t) one-tail

9.92083E-32

P(T<=t) one-tail

0.008153304

t CritiCamphorl one-tail

1.653131676

t CritiCamphorl one-tail

1.653315849

t CritiCamphorl one-tail

1.653975232

P(T<=t) two-tail

9.23057E-35

P(T<=t) two-tail

1.98417E-31

P(T<=t) two-tail

0.016306608

t CritiCamphorl two-tail

1.974185579

t CritiCamphorl two-tail

1.97287136

t CritiCamphorl two-tail

P(T<=t) two tail is

9.23057E-35

1.97315785

P(T<=t) two tail is

1.98417E-31

value < 0.05 - null hypothesis (Ho) rejected

-2.426410306

P(T<=t) two tail is

0.016306608

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Camphor

*Sig difference exists in shoot length between Cineole and Camphor

* Mean shoot length sig decreased in Camphor below Control

* Mean shoot length sig increased in Camphor above Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-60

* Sig difference exists in shoot length between Control and Cineole


* Mean shoot length sig decreased in Cineole below Control

Length
*Pennisetum clandestinum - Radical
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean difference
df

Cineole

Control

16.08

13.74712644

Mean

60.80161616

71.19112537

Variance

100

87

65.63133395

Observations
Pooled Variance

Hypothesized Mean difference

185

df

t Stat

1.964148904

t Stat

P(T<=t) one-tail

0.025505395

P(T<=t) one-tail

t CritiCamphorl one-tail

1.653131676

t CritiCamphorl one-tail

P(T<=t) two-tail

0.051010791

P(T<=t) two-tail

t CritiCamphorl two-tail

P(T<=t) two tail is

1.97287136

t CritiCamphorl two-tail

0.051010791

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Cineole

P(T<=t) two tail is

Camphor

Cineole

Camphor

16.08

19.13414634

Mean

13.74712644

19.13414634

60.80161616

43.94474556

Variance

71.19112537

43.94474556

100

82

87

82

53.21602439

Observations
Pooled Variance

Hypothesized Mean difference

180

df

-2.810217032

t Stat

0.00274941

57.9758154
0
167
-4.596723147

P(T<=t) one-tail

4.21612E-06

1.653363597

t CritiCamphorl one-tail

1.654029802

0.005498821

P(T<=t) two-tail

8.43224E-06

t CritiCamphorl two-tail

1.974271981

1.97323061

0.005498821

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

8.43224E-06

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Camphor

*Sig difference exists in rad. length between Cineole and Camphor

* Mean rad length sig increased in Camphor compared to Control

* Mean rad length sig increased in Camphor compared to Cineole

*Pennisetum clandestinum - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Control

2.23

1.988505747

Mean

0.199090909

0.104517509

Variance

100

87

Observations
Pooled Variance

Cineole

0.155127058

Hypothesized Mean difference

Cineole

Camphor

1.987951807

Mean

1.988505747

1.987951807

0.199090909

0.012048193

Variance

0.104517509

0.012048193

100

83

87

83

0.114353325

Hypothesized Mean difference

185

Camphor
2.23

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

Pooled Variance

df

Observations

0.059383676

Hypothesized Mean difference

181

df

168

t Stat

4.182171563

t Stat

4.820487349

t Stat

0.014815057

P(T<=t) one-tail

2.22669E-05

P(T<=t) one-tail

1.51119E-06

P(T<=t) one-tail

0.494098653

t CritiCamphorl one-tail

1.653131676

t CritiCamphorl one-tail

1.653315849

t CritiCamphorl one-tail

1.653975232

P(T<=t) two-tail

4.45338E-05

P(T<=t) two-tail

3.02238E-06

P(T<=t) two-tail

0.988197306

t CritiCamphorl two-tail

1.974185579

t CritiCamphorl two-tail

1.97287136

t CritiCamphorl two-tail

P(T<=t) two tail is

4.45338E-05

1.97315785

P(T<=t) two tail is

3.02238E-06

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.988197306

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf No. between Control and Camphor


* Mean leaf no. sig reduced in Camphor below Control

* No Sig difference exists in leaf No. between Cineole and Camphor

A11-61

* Sig difference exists in leaf No. between Control and Cineole


* Mean leaf no. sig reduced in Cineole below Control

value > 0.05 - null hypothesis (Ho) accepted

*Pennisetum clandestinum - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

44.56626506

41.17241379

Mean

44.56626506

44.84931507

Mean

41.17241379

44.84931507

Variance

54.51689686

102.5629511

Variance

54.51689686

52.74086758

Variance

102.5629511

52.74086758

83

87

83

73

87

73

Observations
Pooled Variance
Hypothesized Mean difference
df

79.11190081

Observations
Pooled Variance

Hypothesized Mean difference

168

df

53.68654551

Observations
Pooled Variance

Hypothesized Mean difference

154

df

2.486830058

t Stat

P(T<=t) one-tail

0.006932617

P(T<=t) one-tail

0.40503405

P(T<=t) one-tail

0.005214297

t CritiCamphorl one-tail

1.653975232

t CritiCamphorl one-tail

1.65480742

t CritiCamphorl one-tail

1.654555035

P(T<=t) two-tail

0.013865234

P(T<=t) two-tail

1.974185579

t CritiCamphorl two-tail

P(T<=t) two tail is

0.013865234

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Control and Cineole


* Mean leaf area sig reduced in Cineole compared to Control

P(T<=t) two tail is

t Stat

0
158

t Stat

t CritiCamphorl two-tail

-0.240751288

79.85921683

0.8100681
1.975486157

0.8100681

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Camphor

-2.592271654

P(T<=t) two-tail

0.010428593

t CritiCamphorl two-tail

1.975090527

P(T<=t) two tail is

0.010428593

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in leaf area between Cineole and Camphor


* Mean leaf area sig reduced in Cineole compared to Camphor

Pratia purpurascens - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

35.65432099

23.95384615

Mean

35.65432099

16.82258065

Mean

23.95384615

16.82258065

Variance

448.6040123

309.5447115

Variance

448.6040123

102.2466949

Variance

309.5447115

102.2466949

81

65

81

62

65

62

Observations
Pooled Variance

386.7998787

Hypothesized Mean difference

Pooled Variance

df

Observations

298.7614849

Hypothesized Mean difference

144

Pooled Variance

df

Observations

208.3832794

Hypothesized Mean difference

141

df

125

t Stat

3.572589209

t Stat

6.456519814

t Stat

2.782824605

P(T<=t) one-tail

0.000240617

P(T<=t) one-tail

8.05878E-10

P(T<=t) one-tail

0.003112736

t CritiCamphorl one-tail

1.655503183

t CritiCamphorl one-tail

1.655732831

t CritiCamphorl one-tail

1.657135726

P(T<=t) two-tail

0.000481233

P(T<=t) two-tail

1.61176E-09

P(T<=t) two-tail

0.006225472

t CritiCamphorl two-tail

1.976577551

t CritiCamphorl two-tail

1.976932253

t CritiCamphorl two-tail

1.979124136

P(T<=t) two tail is

0.000481233

P(T<=t) two tail is

1.61176E-09

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.006225472

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Camphor

*Sig difference exists in shoot length between Cineole and Camphor

* Mean shoot length sig decreased in Cineole below Control

* Mean shoot length sig decreased in Camphor below Control

* Mean shoot length sig decreased in Camphor below Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-62

* Sig difference exists in shoot length between Control and Cineole

Length
Radicle
Length
Pratia purpurascens - Radical
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

24.15384615

26.10769231

Mean

24.15384615

26.12903226

Mean

26.10769231

26.12903226

Variance

193.2205128

167.8788462

Variance

193.2205128

135.5896351

Variance

167.8788462

135.5896351

91

65

91

62

65

62

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat
P(T<=t) one-tail
t CritiCamphorl one-tail

182.6889111

Observations
Pooled Variance

Hypothesized Mean difference

154

df

-0.890121613

t Stat

0.187394512
1.65480742

169.9391649

Observations
Pooled Variance

Hypothesized Mean difference

151

df

-0.920093806

t Stat

152.1217112
0
125
-0.009746495

P(T<=t) one-tail

0.179495644

P(T<=t) one-tail

0.496119542

t CritiCamphorl one-tail

1.655007509

t CritiCamphorl one-tail

1.657135726

P(T<=t) two-tail

0.374789025

P(T<=t) two-tail

0.358991289

P(T<=t) two-tail

0.992239083

t CritiCamphorl two-tail

1.975486157

t CritiCamphorl two-tail

1.975799933

t CritiCamphorl two-tail

1.979124136

P(T<=t) two tail is

0.374789025

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Cineole

P(T<=t) two tail is

0.358991289

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in rad. length between Control and Camphor

P(T<=t) two tail is

0.992239083

value > 0.05 - null hypothesis (Ho) accepted

*No Sig difference exists in rad. length between Cineole and Camphor

Pratia purpurascens - Leaf Number


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

4.276923077

Mean

4.222222222

5.953365385

Variance

91

65

4.941658342

Hypothesized Mean difference

t Stat

t CritiCamphorl one-tail

4.276923077

3.596774194

4.222222222

3.359333686

Variance

5.953365385

3.359333686

91

62

65

62

3.873638112

P(T<=t) one-tail

1.65480742

Observations
Pooled Variance

4.687477916

Hypothesized Mean difference

151

t Stat

0.222105638

Camphor

Mean

df

-0.767075066

Cineole

3.596774194

Hypothesized Mean difference

154

Camphor
4

Observations
Pooled Variance

df

P(T<=t) one-tail

Control

Observations
Pooled Variance

Cineole

t-Test: Two-Sample Assuming Equal Variances

df

1.24411241
0.107692981

125

t Stat

1.769640442

P(T<=t) one-tail

0.039612712

t CritiCamphorl one-tail

1.655007509

t CritiCamphorl one-tail

1.657135726

P(T<=t) two-tail

0.444211276

P(T<=t) two-tail

0.215385963

P(T<=t) two-tail

0.079225425

t CritiCamphorl two-tail

1.975486157

t CritiCamphorl two-tail

1.975799933

t CritiCamphorl two-tail

1.979124136

P(T<=t) two tail is

0.444211276

P(T<=t) two tail is

0.215385963

value > 0.05 - null hypothesis (Ho) accepted

0.079225425

value > 0.05 - null hypothesis (Ho) accepted

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf No. between Control and Camphor

* No Sig difference exists in leaf No. between Cineole and Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-63

* No Sig difference exists in leaf No. between Control and Cineole

P(T<=t) two tail is

Pratia purpurascens - Leaf Area


t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Camphor

Mean

17.31868132

14.69230769

Mean

17.31868132

13.90322581

Mean

14.69230769

13.90322581

Variance

214.9973138

114.0288462

Variance

214.9973138

130.4167107

Variance

114.0288462

130.4167107

91

65

91

62

65

62

Observations
Pooled Variance
Hypothesized Mean difference
df

173.0363922

Observations
Pooled Variance

Hypothesized Mean difference

154

df

180.8289907

Observations
Pooled Variance

Hypothesized Mean difference

151

df

122.0261241
0
125

t Stat

1.229427316

t Stat

1.542358848

t Stat

0.402389211

P(T<=t) one-tail

0.110393355

P(T<=t) one-tail

0.062539757

P(T<=t) one-tail

0.344042443

t CritiCamphorl one-tail

1.655007509

t CritiCamphorl one-tail

1.657135726

P(T<=t) two-tail

t CritiCamphorl one-tail

0.220786711

1.65480742

P(T<=t) two-tail

0.125079513

P(T<=t) two-tail

0.688084886

t CritiCamphorl two-tail

1.975486157

t CritiCamphorl two-tail

1.975799933

t CritiCamphorl two-tail

1.979124136

P(T<=t) two tail is

0.220786711

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Cineole

P(T<=t) two tail is

0.125079513

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Control and Camphor

P(T<=t) two tail is

0.688084886

value > 0.05 - null hypothesis (Ho) accepted

* No Sig difference exists in leaf area between Cineole and Camphor

Solanum capsicoides - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

24.92307692

23.20253165

Mean

24.92307692

26.89189189

Mean

23.20253165

26.89189189

Variance

15.33846154

37.16358325

Variance

15.33846154

25.57719363

Variance

37.16358325

25.57719363

91

79

91

74

79

74

Observations
Pooled Variance

25.47155376

Hypothesized Mean difference

Pooled Variance

df

19.92390597

Hypothesized Mean difference

168

t Stat

Observations

2.21691054

t Stat

Pooled Variance

df

Observations

31.56221608

Hypothesized Mean difference

163

df

-2.817813496

t Stat

151
-4.059298699

P(T<=t) one-tail

0.013985808

P(T<=t) one-tail

0.002716902

P(T<=t) one-tail

3.93752E-05

t CritiCamphorl one-tail

1.653975232

t CritiCamphorl one-tail

1.654254902

t CritiCamphorl one-tail

1.655007509

P(T<=t) two-tail

0.027971616

P(T<=t) two-tail

0.005433804

P(T<=t) two-tail

7.87504E-05

t CritiCamphorl two-tail

1.974185579

t CritiCamphorl two-tail

1.974622137

t CritiCamphorl two-tail

1.975799933

P(T<=t) two tail is

0.027971616

P(T<=t) two tail is

0.005433804

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

7.87504E-05

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif in shoot length exists between Control and Camphor

* Sig dif in shhot length exists between Cineole and Camphor

* Mean shoot length reduced in Cineole

* Mean shoot length increased in Camphor

* Mean shoot length reduced in Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-64

* Sig dif in shoot length exists between Control and Cineole

Length
Radicle
Length
Solanum capsicoides - Radical
t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Control

35.06329114

Mean

157.3826618

237.0856865

Variance

91

79

Observations
Pooled Variance

Cineole

37.8021978

194.3876375

Hypothesized Mean difference

df

Camphor

Mean

35.06329114

40.94594595

157.3826618

189.5038874

Variance

237.0856865

189.5038874

91

74

79

74

171.7682414

Hypothesized Mean difference

168

Cineole

40.94594595

Observations
Pooled Variance

Camphor

37.8021978

Pooled Variance

df

Observations

214.0825651

Hypothesized Mean difference

163

df

t Stat

1.277475832

t Stat

P(T<=t) one-tail

0.101598427

P(T<=t) one-tail

0.063681708

P(T<=t) one-tail

0.007018655

t CritiCamphorl one-tail

1.653975232

t CritiCamphorl one-tail

1.654254902

t CritiCamphorl one-tail

1.655007509

P(T<=t) two-tail

0.203196855

P(T<=t) two-tail

0.127363416

P(T<=t) two-tail

t CritiCamphorl two-tail

1.974185579

t CritiCamphorl two-tail

1.974622137

t CritiCamphorl two-tail

P(T<=t) two tail is

0.203196855

-1.532395596

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif in rad length between Control and Cineole

t Stat

151

0.127363416

-2.485228956

0.01403731
1.975799933

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif in rad length between Control and Camphor

0.01403731

value < 0.05 - null hypothesis (Ho) rejected

* Cineole sig reduces rad length below Camphor

Solanum capsicoides - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

1.086956522

0.924050633

Mean

1.086956522

0.915492958

Mean

0.924050633

0.915492958

Variance

0.102245581

0.096721844

Variance

0.102245581

0.078470825

Variance

0.096721844

0.078470825

92

79

92

71

79

71

Observations
Pooled Variance

0.099696163

Hypothesized Mean difference

Pooled Variance

df

0.09190873

Hypothesized Mean difference

169

t Stat

Observations

3.3636203

Pooled Variance

df

Observations

0.088089605

Hypothesized Mean difference

161

df

148

t Stat

3.580327459

t Stat

0.176315526

P(T<=t) one-tail

0.000475877

P(T<=t) one-tail

0.000226965

P(T<=t) one-tail

0.430143549

t CritiCamphorl one-tail

1.653920663

t CritiCamphorl one-tail

1.654373136

t CritiCamphorl one-tail

1.655214419

P(T<=t) two-tail

0.000951754

P(T<=t) two-tail

P(T<=t) two-tail

0.860287098

t CritiCamphorl two-tail

1.974099177

t CritiCamphorl two-tail

t CritiCamphorl two-tail

1.976122803

P(T<=t) two tail is

0.000951754

0.00045393
1.974808583

P(T<=t) two tail is

0.00045393

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.860287098

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in leaf No between Control and Camphor


* Mean leaf No reduced by Camphor compared to Control

* No sig dif exists between Cineole and Camphor

A11-65

* Sig dif exists in leaf No between Control and Cineole


* Mean leaf No reduced by Cineole compared to Control

value > 0.05 - null hypothesis (Ho) accepted

Solanum capsicoides - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

6.065217391

2.784810127

Mean

6.065217391

3.27027027

Variance

5.446249403

4.504381694

Variance

5.446249403

4.583487597

92

79

92

74

Observations
Pooled Variance

5.01154123

Hypothesized Mean difference

df
t Stat

Observations
Pooled Variance

5.062215184

Hypothesized Mean difference

169

df

9.55325819

2.784810127

3.27027027

Variance

4.504381694

4.583487597

79

74

Observations
Pooled Variance

Hypothesized Mean difference

164

Camphor

Mean

df

4.542624945
0
151

t Stat

7.955345103

t Stat

P(T<=t) one-tail

7.29671E-18

P(T<=t) one-tail

1.38649E-13

P(T<=t) one-tail

0.080602281

t CritiCamphorl one-tail

1.653920663

t CritiCamphorl one-tail

1.654198059

t CritiCamphorl one-tail

1.655007509

P(T<=t) two-tail

1.45934E-17

P(T<=t) two-tail

2.77298E-13

P(T<=t) two-tail

0.161204561

t CritiCamphorl two-tail

1.974099177

t CritiCamphorl two-tail

1.974535735

t CritiCamphorl two-tail

1.975799933

P(T<=t) two tail is

1.45934E-17

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

2.77298E-13

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists in leaf area between Control and Cineole

* Sig dif exists in leaf area between Control and Camphor

* Mean leaf area reduced by Cineole

* Mean leaf area reduced by Camphor

-1.407938969

0.161204561

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif between Cineole and Camphor

Syzygium leuhmannii - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

51.34782609

43.13829787

Mean

51.34782609

51

Variance

81.52603918

79.77636696

Variance

81.52603918

84.68817204

92

94

92

94

Observations
Pooled Variance

80.64169398

Hypothesized Mean difference

Pooled Variance

df

Observations

83.12429112

Hypothesized Mean difference

184

43.13829787

51

Variance

79.77636696

84.68817204

94

94

Observations
Pooled Variance

df

Camphor

Mean

82.2322695

Hypothesized Mean difference

184

df

186

t Stat

6.233611716

t Stat

0.260135451

t Stat

P(T<=t) one-tail

1.51848E-09

P(T<=t) one-tail

0.397525079

P(T<=t) one-tail

6.75142E-09

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653088475

P(T<=t) two-tail

3.03696E-09

P(T<=t) two-tail

0.795050158

P(T<=t) two-tail

1.35028E-08

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

P(T<=t) two tail is

3.03696E-09

P(T<=t) two tail is

0.795050158

value < 0.05 - null hypothesis (Ho) rejected

*Sig.dif exists between Control and Cineole

-5.943528856

1.9727986

P(T<=t) two tail is

1.35028E-08

value > 0.05 - null hypothesis (Ho) accepted

* Not sig. dif exists between Control and Camphor

value < 0.05 - null hypothesis (Ho) rejected

* Sig. dif exists between Cineole and Camphor


* Cineole is reduced below Camphor

A11-66

* Cineole is reduced below Control

Length
Syzygium leuhmannii - Radical
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Control

52.61956522

43.26595745

101.908624

64.7994738

92

94

Observations
Pooled Variance

Cineole

83.15236873

Hypothesized Mean difference

Mean
Variance

Camphor
48.74468085

Mean

101.908624

125.5470144

Variance

92

94

113.8562887

Hypothesized Mean difference

184

Cineole

52.61956522

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

48.74468085

64.7994738

125.5470144

94

94

Observations
Pooled Variance

df

Camphor

43.26595745

95.17324411

Hypothesized Mean difference

184

df

0
186

t Stat

6.994283154

t Stat

2.476176534

t Stat

P(T<=t) one-tail

2.39115E-11

P(T<=t) one-tail

0.007091656

P(T<=t) one-tail

8.11393E-05

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653088475

P(T<=t) two-tail

4.78229E-11

P(T<=t) two-tail

0.014183313

P(T<=t) two-tail

0.000162279

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

P(T<=t) two tail is

4.78229E-11

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.014183313

-3.850089877

1.9727986

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.000162279

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists between Control and Cineole rad length

* Sig dif exists between Control and Camphor rad length

* Sig dif exists between Cineole and Camphor rad length

* Mean rad length Cineole is reduced below Control

* Mean rad length Camphor is reduced below Control

* Mean rad length Cineole is reduced below Camphor

Syzygium leuhmannii - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Camphor

Mean

5.315217391

4.553191489

Mean

5.315217391

4.989361702

Mean

4.553191489

4.989361702

Variance

1.075370282

1.647677877

Variance

1.075370282

2.010638298

Variance

1.647677877

2.010638298

92

94

92

94

94

94

Observations
Pooled Variance

1.364634447

Hypothesized Mean difference

df

1.548087268

Hypothesized Mean difference

184

t Stat

Observations
Pooled Variance

df

4.44798114

1.829158087

Hypothesized Mean difference

184

t Stat

Observations
Pooled Variance

df

1.78578412

t Stat

186
-2.210951986

P(T<=t) one-tail

7.48682E-06

P(T<=t) one-tail

0.037890566

P(T<=t) one-tail

0.014129049

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653088475
0.028258098

P(T<=t) two-tail

1.49736E-05

P(T<=t) two-tail

0.075781132

P(T<=t) two-tail

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

P(T<=t) two tail is

1.49736E-05

P(T<=t) two tail is

0.075781132

value < 0.05 - null hypothesis (Ho) rejected

*Sig dif exists for leaf No between Control and Cineole

1.9727986

P(T<=t) two tail is

0.028258098

value > 0.05 - null hypothesis (Ho) accepted

* No sig dif exists in leaf No between Control and Camphor

value < 0.05 - null hypothesis (Ho) rejected

* Sig dif exists between Cineole and Camphor leaf no


* Cineole leaf no is reduced below Camphor

A11-67

*Mean leaf no reduced in Cineole compared to Control

Syzygium leuhmannii - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

87.34782609

80.76595745

Variance

955.2183469

922.912377

92

94

Observations
Pooled Variance

938.889786

Hypothesized Mean difference


df

t-Test: Two-Sample Assuming Equal Variances

Cineole

87.34782609

84.54255319

Mean

Variance

955.2183469

1119.756234

Variance

92

94

Observations
Pooled Variance
Hypothesized Mean difference

184

Camphor

Mean

df

1038.381518

Observations
Pooled Variance

Hypothesized Mean difference

184

df

Camphor

80.76595745

84.54255319

922.912377

1119.756234

94

94

1021.334306
0
186

t Stat

1.464681754

t Stat

0.593605627

t Stat

-0.81015006

P(T<=t) one-tail

0.072357139

P(T<=t) one-tail

0.276752467

P(T<=t) one-tail

0.209444613

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653088475

P(T<=t) two-tail

0.144714279

P(T<=t) two-tail

0.553504934

P(T<=t) two-tail

0.418889226

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

P(T<=t) two tail is

0.144714279

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

*No sig dif exists in leaf area between Control and Cineole

0.553504934

value > 0.05 - null hypothesis (Ho) accepted

*No sig dif exists in leaf area between Control and Camphor

1.9727986

P(T<=t) two tail is

0.418889226

value > 0.05 - null hypothesis (Ho) accepted

*No sig dif exists in leaf area between Cineole and Camphor

Syzygium paniculatum - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

43.23913043

37.65116279

Variance

140.2718586

132.276881

92

86

Observations
Pooled Variance

136.4106478

Hypothesized Mean difference

Camphor

43.23913043 31.70212766

Mean

Variance

140.2718586 187.9748341

Variance

Observations

92

94

164.3826017

Hypothesized Mean difference

176

Cineole

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

df

31.70212766

132.276881

187.9748341

86

94

Observations
Pooled Variance

161.377497

Hypothesized Mean difference

184

Camphor

37.65116279

df

178

t Stat

3.189795825

t Stat

6.135732828

t Stat

3.138354118

P(T<=t) one-tail

0.000842894

P(T<=t) one-tail

2.53766E-09

P(T<=t) one-tail

0.000994502

t CritiCamphorl one-tail

1.653556865

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653459094

P(T<=t) two-tail

0.001685787

P(T<=t) two-tail

5.07533E-09

P(T<=t) two-tail

0.001989005

t CritiCamphorl two-tail

1.973535291

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

1.973380677

P(T<=t) two tail is

0.001685787

P(T<=t) two tail is

5.07533E-09

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.001989005

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* A significant difference exists in the shoot length between the Control and Cineole treatments

* significant difference exists in the shoot length between Control and Camphor treatments

* A significant difference exists in the shoot length between the Cineole and Camphor treatments

* Mean shoot length is reduced in the Cineole treatment

* Mean shoot length is reduced in the Camphor treatment

* Camphor mean shoot length is less than Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Syzygium paniculatum - Radical


Length
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

38.09782609

32

Variance

90.57274247

104.2352941

92

86

Observations
Pooled Variance

97.17113389

Hypothesized Mean difference

t Stat

Mean

Variance

90.57274247 177.9720888

Variance

Observations

92

Observations
Pooled Variance

df

4.12419828

94

134.7474121

Hypothesized Mean difference

176

Cineole

38.09782609 29.53191489

Pooled Variance

df

Camphor

Mean

Hypothesized Mean difference

184

df

Camphor
32

29.53191489

104.2352941

177.9720888

86

94

142.7606981
0
178

t Stat

5.031699765

t Stat

1.384307908

P(T<=t) one-tail

2.86358E-05

P(T<=t) one-tail

5.75514E-07

P(T<=t) one-tail

0.083998609

t CritiCamphorl one-tail

1.653556865

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653459094

P(T<=t) two-tail

5.72715E-05

P(T<=t) two-tail

1.15103E-06

P(T<=t) two-tail

0.167997219

t CritiCamphorl two-tail

1.973535291

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

1.973380677

P(T<=t) two tail is

5.72715E-05

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

1.15103E-06

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

A11-68

t-Test: Two-Sample Assuming Equal Variances

0.167997219

value > 0.05 - null hypothesis (Ho) not rejected

* A significant difference. in rad. length exists between the Control and Cineole treatments

* A significant difference. in rad. length exists between the Control and Camphor treatments

* Not sig.difference in rad length exists between the Cineole and Camphor treatments

* Mean rad. length reduced in Cineole treatment

* Mean rad. length reduced in Camphor treatment

* Mean rad. length similar

Syzygium paniculatum - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Mean

3.760869565

3.11627907

Variance

3.766364071

3.256908345

92

86

Observations
Pooled Variance

3.520320112

Hypothesized Mean difference

Cineole

3.760869565 1.957446809

Mean

Variance

3.766364071 4.213223519

Variance

Observations

92

94

3.992222379

Hypothesized Mean difference

176

Camphor

Mean

Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

df

1.957446809

3.256908345

4.213223519

86

94

Observations
Pooled Variance

3.75655616

Hypothesized Mean difference

184

Camphor

3.11627907

df

178

t Stat

2.290477117

t Stat

6.154482648

t Stat

P(T<=t) one-tail

0.011589874

P(T<=t) one-tail

2.30079E-09

P(T<=t) one-tail

4.51656E-05

t CritiCamphorl one-tail

1.653556865

t CritiCamphorl one-tail

1.653177151

t CritiCamphorl one-tail

1.653459094

P(T<=t) two-tail

0.023179747

P(T<=t) two-tail

4.60159E-09

P(T<=t) two-tail

9.03311E-05

t CritiCamphorl two-tail

1.973535291

t CritiCamphorl two-tail

1.972939572

t CritiCamphorl two-tail

1.973380677

P(T<=t) two tail is

0.023179747

P(T<=t) two tail is

4.60159E-09

value < 0.05 - null hypothesis (Ho) rejected

4.006844922

P(T<=t) two tail is

9.03311E-05

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

*Sig.difference exists in leaf No. between the Control and Camphor treatments

*Sig.difference exists in leaf No. between the Cineole and Camphor treatments

*Mean leaf No. is reduced in Camphor treatment

*Mean leaf No. is reduced in Camphor treatment

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-69

*Sig.difference exists in leaf No. between the Control and Cineole treatments
*Mean leaf No. is reduced in Cineole treatment

Syzygium paniculatum - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean difference
df

Cineole

Control

31.67391304

24.84883721

Mean

821.10129

591.5886457

Variance

92

86

710.2571152
0

92

85

596.5316624

Hypothesized Mean difference

176

Cineole

821.10129 353.2478992

Observations
Pooled Variance

Camphor

31.67391304 16.11764706

24.84883721

16.11764706

Variance

591.5886457

353.2478992

86

85

Observations
Pooled Variance

df

Hypothesized Mean difference

175

Camphor

Mean

df

473.1234226
0
169

t Stat

1.707390351

t Stat

4.233556436

t Stat

2.624501682

P(T<=t) one-tail

0.044756457

P(T<=t) one-tail

1.85417E-05

P(T<=t) one-tail

0.004736343

t CritiCamphorl one-tail

1.653556865

t CritiCamphorl one-tail

1.653606887

t CritiCamphorl one-tail

1.653920663

P(T<=t) two-tail

0.089512914

P(T<=t) two-tail

3.70834E-05

P(T<=t) two-tail

0.009472686

t CritiCamphorl two-tail

1.973535291

t CritiCamphorl two-tail

1.973612598

t CritiCamphorl two-tail

1.974099177

P(T<=t) two tail is

0.089512914

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) not rejected

* No sig. difference. in leaf area between Control and Cineole

3.70834E-05

value < 0.05 - null hypothesis (Ho) rejected

P(T<=t) two tail is

0.009472686

value < 0.05 - null hypothesis (Ho) rejected

* Sig.difference in leaf area between Control and Camphor treatments

* Sig.difference in leaf area between Cineole and Camphor treatments

* Mean leaf area reduced in Camphor treatment

* Mean leaf area is sig. reduced by Camphor between Cineole And Camphor treatments

Toona australis - Shoot Length


t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

21.78947368

23.38095238

Mean

21.78947368

Variance

27.12842105

34.09409065

Variance

27.12842105 50.44816468

76

84

Observations
Pooled Variance

30.78760192

Hypothesized Mean difference

df

Observations
Pooled Variance

76

t Stat

-1.81175449

t Stat

0.035961566

P(T<=t) one-tail

t CritiCamphorl one-tail

1.654555035

t CritiCamphorl one-tail

P(T<=t) two-tail

0.071923133

t CritiCamphorl two-tail

1.975090527

P(T<=t) two tail is

0.071923133

23.38095238

25.109375

Variance

34.09409065

50.44816468

84

64

Observations

41.15098561

Hypothesized Mean difference

138

df

-3.183898426

t Stat

0.00089808

Camphor

Mean

Pooled Variance

df

P(T<=t) one-tail

64

37.77439097

Hypothesized Mean difference

158

25.109375

146
-1.623897423

P(T<=t) one-tail

0.053277387

1.655971573

t CritiCamphorl one-tail

1.655357664

P(T<=t) two-tail

0.001796161

P(T<=t) two-tail

0.106554774

t CritiCamphorl two-tail

1.977305146

t CritiCamphorl two-tail

1.976345629

P(T<=t) two tail is

0.001796161

value > 0.05 - null hypothesis (Ho) accepted

P(T<=t) two tail is

0.106554774

value < 0.05 - null hypothesis (Ho) rejected

value > 0.05 - null hypothesis (Ho) accepted

* No sig.dif between shoot lengths Control and Cineole treatments

* Sig.dif in shoot length between Control and Camphor treatments

* No sig.dif in shoot length between Cineole and Camphor treatments

* Mean shoot length similar

* Mean shoot length increased by Camphor treatment

* Mean shoot length similar between Cineole and Camphor treatments

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-70

t-Test: Two-Sample Assuming Equal Variances

Toona australis - Radical


Radicle
Length
Length
t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance

Control

14.82142857

Mean

80.18666667

59.81110155

Variance

76

84

Observations
Pooled Variance

Cineole
19

69.48304702

Hypothesized Mean difference

df

76

64

73.5234375

Hypothesized Mean difference

158

Cineole

13.109375

80.18666667 65.59102183

Observations
Pooled Variance

Camphor
19

14.82142857

13.109375

Variance

59.81110155

65.59102183

84

64

Observations
Pooled Variance

df

Hypothesized Mean difference

138

Camphor

Mean

df

62.30517674
0
146

t Stat

3.166466157

t Stat

4.049311992

t Stat

P(T<=t) one-tail

0.000926033

P(T<=t) one-tail

4.26244E-05

P(T<=t) one-tail

t CritiCamphorl one-tail

1.654555035

t CritiCamphorl one-tail

1.655971573

t CritiCamphorl one-tail

1.655357664

P(T<=t) two-tail

0.001852065

P(T<=t) two-tail

8.52488E-05

P(T<=t) two-tail

0.193187521

t CritiCamphorl two-tail

1.975090527

t CritiCamphorl two-tail

1.977305146

t CritiCamphorl two-tail

1.976345629

P(T<=t) two tail is

0.001852065

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

8.52488E-05

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

1.307236171
0.09659376

0.193187521

value > 0.05 - null hypothesis (Ho) accepted

* Sg.dif in rad. length exists between the Control and Cineole treatments.

* Sg.dif in rad. length exists between the Control and Camphor treatments.

*No Sg.dif in rad. length exists between the Cineole and Camphor treatments.

* Mean Rad. length reduced in Cineole treatment

* Mean Rad. length reduced in Camphor treatment

* Mean Rad. length is similar

Toona australis - Leaf Number

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

t-Test: Two-Sample Assuming Equal Variances

Control

Camphor

Cineole

Mean

1.144736842

1.540540541

Mean

1.144736842

0.90625

Variance

0.632105263

0.553128471

Variance

0.632105263

0.75297619

76

74

76

64

Observations
Pooled Variance

0.593150494

Hypothesized Mean difference

t Stat

Pooled Variance

df

Observations

0.687285469

Hypothesized Mean difference

148
-3.146838292

1.540540541

0.90625

Variance

0.553128471

0.75297619

74

64

Observations
Pooled Variance

df

0.645704988

Hypothesized Mean difference

138

Camphor

Mean

df

136

t Stat

1.6956203

t Stat

P(T<=t) one-tail

0.000997993

P(T<=t) one-tail

0.0461064

P(T<=t) one-tail

4.32619E-06

t CritiCamphorl one-tail

1.655214419

t CritiCamphorl one-tail

1.655971573

t CritiCamphorl one-tail

1.656135282

P(T<=t) two-tail

0.001995986

P(T<=t) two-tail

0.092212799

P(T<=t) two-tail

8.65238E-06

t CritiCamphorl two-tail

1.976122803

t CritiCamphorl two-tail

1.977305146

t CritiCamphorl two-tail

1.977559805

P(T<=t) two tail is

0.001995986

P(T<=t) two tail is

0.092212799

value < 0.05 - null hypothesis (Ho) rejected

4.624209138

P(T<=t) two tail is

8.65238E-06

value > 0.05 - null hypothesis (Ho) accepted

value < 0.05 - null hypothesis (Ho) rejected

* Sg.dif in leaf.no. exists between the Control and Cineole treatments.

* No sig.dif in leaf no exists between the Control and Camphor treatments.

* Sg.dif in leaf no. exists between the Cineole and Camphor treatments.

* Mean leaf no. increased in Cineole treatment

* Mean leaf no. is similar

* Mean leaf no. is reduced in Camphor

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-71

t-Test: Two-Sample Assuming Equal Variances

Toona australis - Leaf Area

t-Test: Two-Sample Assuming Equal Variances

Control

Cineole

Control

Camphor

Cineole

Mean

8.697368421

11.11904762

Mean

8.697368421

Variance

283.4938596

182.0097533

Variance

283.4938596 49.64260913

76

84

Observations
Pooled Variance
Hypothesized Mean difference
df
t Stat

230.1825886

Observations
Pooled Variance

76

11.11904762

4.234375

Variance

182.0097533

49.64260913

84

64

Observations

124.892424

Hypothesized Mean difference

138

Camphor

Mean

Pooled Variance

df

-1.008246735

64

176.7356801

Hypothesized Mean difference

158

4.234375

df

146

t Stat

1.978777184

t Stat

3.712906548

P(T<=t) one-tail

0.157438803

P(T<=t) one-tail

0.024915927

P(T<=t) one-tail

0.000145426

t CritiCamphorl one-tail

1.654555035

t CritiCamphorl one-tail

1.655971573

t CritiCamphorl one-tail

1.655357664

P(T<=t) two-tail

0.314877605

P(T<=t) two-tail

0.049831855

P(T<=t) two-tail

0.000290851

t CritiCamphorl two-tail

1.975090527

t CritiCamphorl two-tail

1.977305146

t CritiCamphorl two-tail

1.976345629

P(T<=t) two tail is

0.314877605

P(T<=t) two tail is

value > 0.05 - null hypothesis (Ho) accepted

* No sig. difference. in leaf area between Control and Cineole

0.049831855

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.000290851

value < 0.05 - null hypothesis (Ho) rejected

* Sig.difference in leaf area between Control and Camphor treatments

* Sig.difference in leaf area between Cineole and Camphor treatments

* Mean leaf area reduced in Camphor treatment

* Mean leaf area is sig. reduced by Camphor

Viola hederacea - Shoot Length

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

Control
Mean

Control

40.25510204

33.37096774

Mean

136.748685

183.2863564

Variance

98

62

Variance
Observations
Pooled Variance

Cineole

154.7157607

Hypothesized Mean difference

Cineole

Camphor

44.55932203

Mean

33.37096774

44.55932203

136.748685

224.7679719

Variance

183.2863564

224.7679719

98

59

62

59

169.6849343

Hypothesized Mean difference

158

Camphor

40.25510204

Observations
Pooled Variance

df

t-Test: Two-Sample Assuming Equal Variances

Pooled Variance

df

Observations

203.5042867

Hypothesized Mean difference

155

df

t Stat

3.41060125

t Stat

P(T<=t) one-tail

0.00041149

P(T<=t) one-tail

-2.005220037
0.023340058

P(T<=t) one-tail

t Stat

119
-4.312296311
1.67604E-05

t CritiCamphorl one-tail

1.654555035

t CritiCamphorl one-tail

1.654743755

t CritiCamphorl one-tail

P(T<=t) two-tail

0.000822979

P(T<=t) two-tail

0.046680115

P(T<=t) two-tail

3.35208E-05

t CritiCamphorl two-tail

1.975090527

t CritiCamphorl two-tail

1.975386112

t CritiCamphorl two-tail

1.980097295

P(T<=t) two tail is

0.000822979

P(T<=t) two tail is

0.046680115

value < 0.05 - null hypothesis (Ho) rejected

1.65775873

P(T<=t) two tail is

3.35208E-05

value < 0.05 - null hypothesis (Ho) rejected

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in shoot length between Control and Camphor

*Sig difference exists in shoot length between Cineole and Camphor

* Mean shoot length sig decreased in Cineole below Control

* Mean shoot length sig increased in Camphor above Control

* Mean shoot length sig increased in Camphor above Cineole

t-Test: Two-Sample Assuming Equal Variances

t-Test: Two-Sample Assuming Equal Variances

A11-72

* Sig difference exists in shoot length between Control and Cineole

Viola hederacea - Radical


Length
Radicle
Length
t-Test: Two-Sample Assuming Equal Variances

Control
Mean
Variance
Observations
Pooled Variance

Cineole
11.08064516

Mean

152.128866

80.69830777

Variance

98

62

124.5512454

Hypothesized Mean difference


df

Control

18.07142857

Observations
Pooled Variance

Hypothesized Mean difference

158

df

Camphor

Cineole

Camphor

18.07142857

11.11864407

Mean

11.08064516

11.11864407

152.128866

54.03740503

Variance

80.69830777

54.03740503

98

59

62

59

115.4236741

Observations
Pooled Variance

Hypothesized Mean difference

155

df

67.7039182
0
119

t Stat

3.860119068

t Stat

3.927358432

t Stat

P(T<=t) one-tail

8.24248E-05

P(T<=t) one-tail

6.44732E-05

P(T<=t) one-tail

t CritiCamphorl one-tail

1.654555035

t CritiCamphorl one-tail

1.654743755

t CritiCamphorl one-tail

P(T<=t) two-tail

0.000128946

P(T<=t) two-tail

0.979784988

t CritiCamphorl two-tail

1.975386112

t CritiCamphorl two-tail

1.980097295

P(T<=t) two-tail
t CritiCamphorl two-tail

0.00016485
1.975090527

P(T<=t) two tail is

0.00016485

P(T<=t) two tail is

value < 0.05 - null hypothesis (Ho) rejected

0.000128946

value < 0.05 - null hypothesis (Ho) rejected

* Sig difference exists in rad. length between Control and Cineole

* Sig difference exists in rad. length between Control and Camphor

* Mean rad length sig decreased in Cineole below Control

* Mean rad length sig decreased in Camphor below Control

P(T<=t) two tail is

-0.025391793
0.489892494
1.65775873

0.979784988

value > 0.05 - null hypothesis (Ho) accepted

*No Sig difference exists in rad. length between Cineole and Camphor

__________________________________________________________________________________________________________________________________________________________________________________
1
Note: Some species were not tested due to very low germination (*Lantana camara and Geitonoplesium cymosum); missing data (Ficus macrophylla, Eucalyptus microcorys, *Setaria sphaecelata, Ottochloa gracilima and
*Chloris gayana) or zero values recorded for germination (Mallotus philippensis).

Appendix 12.

Summary Effect Tables

derived from the t-tests of shoot length, radical length, leaf number and leaf area
for all species tested.
Table A12.1 Summary effect table for t-tests of post-emergent growth in the trees and
shrubs.
Species

Treatment Significance to
Control.

Significance
Between
Treatments.
Mean Increased or
Decreased

Mean Comparison
Cineole to Camphor

Acacia melanoxylon
Ci

Radicle Length

ns

ns

Leaf No.

ns

ns

ns

ns

Ci

>

ns

ns

Leaf No.

ns

ns

Leaf Area

ns

ns

Shoot Length

Leaf Area

Allocasuarina littoralis
Shoot Length
Radicle Length

Allocasuarina torulosa
Ci

Radicle Length

ns

ns

Leaf No.

ns

ns

<

ns

ns

ns

ns

Radicle Length

ns

ns

>

Leaf No.

ns

ns

ns

ns

<

Shoot Length

na3 Ca

na

Radicle Length

na Ca

na3

Leaf No.

na3 Ca

na3

Leaf Area

na3 Ca

na

Ca

<

ns

ns

Shoot Length

Leaf Area
Araucaria cunninghamiana
Shoot Length

Leaf Area

Callistemon viminalis

*Cassia coleutioides
Shoot Length
Radicle Length
Leaf No.
Leaf Area

na

na

na1

na1

na
na

*Cinnamomum camphora
(from soil)
Shoot Length

ns

ns

Radicle Length

Ci

<

Leaf No.

ns

ns

Leaf Area

ns

ns

Shoot Length

Ci Ca

++

Radicle Length

Ci Ca

++

>

Leaf No.

Ci Ca

++

Ci Ca

++

>

ns

ns

ns

ns

Leaf No.

ns

ns

Leaf Area

Ci Ca

--

Ci

<

Ci Ca

--

Leaf No.

ns

ns

Leaf Area

ns

ns

Ci

<

ns

ns

Leaf No.

Ci

<

Leaf Area

Ci Ca

--

*Cinnamomum camphora
(fresh from tree)

Leaf Area

Commersonia bartramia
Shoot Length
Radicle Length

Cupaniopsis parvifolia
Shoot Length
Radicle Length

Eucalyptus intermedia
Shoot Length
Radicle Length

Eucalyptus microcorys

na

na

na

Shoot Length

Ci

Radicle Length

Ca

>

Leaf No.

Ci Ca

--

>

Leaf Area

Ci Ca

--

>

Ci Ca

++

>

Ci Ca

++

>

Eucalyptus saligna

Ficus coronata
Shoot Length
Radicle Length

A12-2

Leaf No.

Ca

>

Leaf Area

Ca

>

Ficus macrophylla

na

na

na

Ci

<

Radicle Length

Ci

<

Leaf No.

ns

ns

Ci Ca

--

<

Shoot Length

Ci

<

Radicle Length

ns

ns

Leaf No.

Ci

<

Leaf Area

Ci

<

na1

na1

na1

Shoot Length

Ci Ca

--

<

Radicle Length

Ci Ca

--

<

Leaf No.

Ci Ca

--

<

Ci

<

Ca

<

Radicle Length

ns

ns

<

Leaf No.

ns

ns

Ca

Shoot Length

Ci Ca

--

Radicle Length

Ci Ca

--

Leaf No.

Ci Ca

--

Leaf Area

Ci Ca

--

Shoot Length

Ci

<

Radicle Length

ns

ns

Leaf No.

Ci

Leaf Area

Ci

na1

na1

na1

Guioa semiglauca
Shoot Length

Leaf Area

Hymenosporum flavum

*Lantana camara

* Ligustrum lucidum

Leaf Area

Lepiderema pulchella
Shoot Length

Leaf Area

Lophostemon confertus

Macaranga tanarius

Mallotus philippensis

A12-3

Omalanthus populifolius
ns

Ci

>

Leaf No.

ns

ns

Leaf Area

Ca

Ci Ca

-+

<

ns

ns

<

Leaf No.

Ci Ca

--

Leaf Area

Ci Ca

--

Ci

<

Ci Ca

--

<

Ci

<

ns

ns

Shoot Length

Ci Ca

--

Radicle Length

Ci Ca

--

Leaf No.

Ci Ca

--

>

Leaf Area

Ca

>

Ca

Ci Ca

--

Leaf No.

Ci

>

Leaf Area

Ca

<

na2

na2

na2

Shoot Length
Radicle Length

Ci Ca

Solanum capsicoides
Shoot Length
Radicle Length

Syzgium luehmannii
Shoot Length
Radicle Length
Leaf No.
Leaf Area

Syzgium paniculatum

Toona australis
Shoot Length
Radicle Length

Tristaniopsis laurina

Key:
Ci : 'Cineole' treatment
Ca : 'Camphor' treatment.
ns : no significant difference.
+ : mean of treatment significantly increased above the 'Control'.
- : mean of treatment significantly reduced below the 'Control'.
- - , - +, etc : first symbol identifies significant effect direction in 'Cineole' t