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2010
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
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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,
ii
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
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
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
9
11
15
41
vii
1.3
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
57
2.1
Introduction
57
2.2
58
2.3
Methods
58
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
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
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
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
149
4.1
Introduction
149
4.2
150
4.3
Methods
151
4.4
Results
153
4.4.1
4.4.2
4.4.3
154
154
163
4.5
Discussion
165
4.6
Conclusion
169
ix
171
5.1
172
5.2
172
174
5.3
REFERENCES
179
LIST OF TABLES
Table No.
1.
2.
3.
4.
5.
6.
7.
Title
Page
14
14
36
42
51
75
77
78
94
10.
99
11.
121
135
8.
9.
12.
xi
13.
14.
15.
141
141
144
LIST OF FIGURES
Figure No.
1.
Title
Page
46
50
52
60
5.
71
6.
72
2.
3.
4.
7.
8.
9.
74
79
96
xii
10.
97
11.
97
12.
100
101
102
102
103
103
103
110
112
114
116
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
xiii
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
116
117
118
118
140
142
142
156
157
159
161
163
xiv
LIST OF PLATES
Plate No.
1.
Title
Page
177
LIST OF APPENDICES
A1
A2
A3
A4
A5
A6
A7
A8
A9
A10
A11
A12
A13
A14
A15
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
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).
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
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.
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
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
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
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
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,
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
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).
12
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.
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.
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
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.
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:
15
ruderals (R), allowing exploitation under conditions of low stress and high
disturbance.
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),
16
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.
18
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.
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).
24
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
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.
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.
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.).
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.
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
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
(Woodford 1993)
(Holmes 1987)
36
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
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).
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
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).
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.
43
How is camphor laurel able to be such an efficient and effective intrusive plant
i.e. what physical and ecological factors are involved?
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
acetate
cinnamic acid
derivatives
mevalonic
acid
courmarins
flavonoids
condensed tannins
terpenoids
and
steroids
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
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).
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
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
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.
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.
review of what is known worldwide about the trees ecological strategies and the
status it has reached as an intrusive exotic plant within Australia;
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;
(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
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
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
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
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.
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.
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).
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.
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.
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
50
shade installed
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
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 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 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
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.
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.
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
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 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;
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
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
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
Herb s
response in
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%.
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%.
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.
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%.
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.
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.
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.
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.
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.
85
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.
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.
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 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.
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.
*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
*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
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%.
92
day 41 at 15% for the control, day 47 at 10% for cineole and day 57 at 15% for
camphor.
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)
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
(b).
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
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.
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
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
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.
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
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.
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
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
1.8
1.6
1.4
1.2
1
Co
Ca
Ci
T re a t m e nt
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
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
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
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
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.
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.
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
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.
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
Both Trea tm en ts
Significant
No Significant
D ifference
(b).
18
No. of species with significant
differences.
18
16
n = 36
14
62+
12
10
8
73+
1Ci-Ca+
13
31+
6
4
2
0
'Cineole' only
No Significant
Difference
109
(c).
16
18
16
n = 35
14
12
10
101+
42+
2-
6
4
2
0
'Cineole' only
'Camphor' only
Both Treatments
Significant
No Significant
Difference
18
16
14
12
10
8
101+
1Ci-C a+
10
931+
6
4
2
0
'Cineole' only
'Camphor' only
Both Treatments
Significant
No Significant
Difference
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.
(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
shrub species
showing
significant
Figure X. Number of tree and
'shoot length' variation in the post-emergent measurements.
(b).
11
14
62+
12
10
n = 24
8
6
21+
2-
2
0
'Cineole' only
'Camphor' only
Both Treatments
Significant
No Significant
Difference
111
(c).
14
12
n = 23
61+
10
8
10
22+
6
2-
4
2
0
'Cineole' only
'Camphor' only
Both Treatments
Significant
No Significant
Difference
(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
variation
in theforpost-emergent
analysis
not available
one species)
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).
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
(b).
14
12
n = 9
10
31+
11 Ci-Ca+
6
4
1-
2
0
'C ineole' only
No Signific ant
D ifferenc e
(c).
14
12
n = 9
10
8
6
4
2
0
32-
2-
'Cineole' only
'Camphor' only
Both Treatments
Significant
No Significant
Difference
113
differences
(d).
14
12
10
8
6
4
2
0
n = 9
31Ci-Ca+
2
21-
'Camphor' on ly
No Significant
Difference
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).
significant differences
14
12
n = 3
10
8
6
1-
11-
4
2
0
B oth Treatments
S ignificant
No S ignificant
D ifference
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).
14
12
n = 3
10
8
6
1-
11+
1- 11+
2
0
'C ineole' only
Both Treatments
Significant
No Significant
Difference
T
re atme nt Significance
SignificancetotoControl
'C ontrol'
Treatment
(c).
14
12
n= 3
10
8
6
1-
4
2
0
'Cineole' only
115
No Significant
Difference
Difference
(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
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).
25
21
21
19
16
20
14
15
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
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.
14
13
13
12
12
11
'C in e o le ' > 'C a m p h o r'
10
10
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
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
10
differences
n = 9
12
8
6
4
5
3
3
2
0
s hoot length
radic al length
leaf No.
leaf area
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
14
12
differences
size as this identified that it was not representative of the vines per se.
n =3
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
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.
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
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.
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
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
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.
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,
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
127
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.
128
2.6
Conclusion
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.
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
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
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.
132
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:
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
Toluidine Blue
lignin, tannins
polyphenols i.e.
pectin, pectic
substances.
Phloroglucinol
lignin
IK Iodine
starch, cellulose
Methylene Blue
cellulose
Ferric Sulphate
tannins
Safranin Red
Sudan III
cutin, chromatin,
lignin, phenolics,
tannins, suberin
chloroplasts,
chitin, nuclei &
chromosomes.
oils, fats, cutin
Red Henna
camphor?
glycogen
calcium oxalate
Pizzolata (1964)
Cochineal
(simple carmine)
Pizzolato
structures in microns (m) was further undertaken where visual differences appeared
worthy of further validation.
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.
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:
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.
These
included
-terpinene,
-terpinolene,
-terpineol,
ledene,
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
globules
cutinized layer
air space
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
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
70
'Cineole' chemo-type
60
50
40
'Camphor' chemo-type
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 97 101 105
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
80
Globule Type 3.
70
60
50
40
Globule Type1.
30
20
10
0
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.
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
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%).
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
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
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.
150
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.
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.
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
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
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
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
120
100
80
60
Control
Camphor
40
20
0
0
200
400
600
800
Number of seedlings measured
1000
1200
(b).
60
n = 895
50
40
n = 677
30
Control
20
Camphor
10
0
0
200
400
600
Number of seedlings measured
800
1000
1200
(c).
5000
4500
4000
3500
n = 895
3000
2500
n = 677
2000
1500
1000
Control
500
Camphor
0
0
200
400
600
800
1000
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
140
120
100
80
60
Control
Camphor
40
20
0
0
200
400
600
800
Number of seedlings measured
158
1000
1200
(b).
60
50
40
30
20
Control
n = 128
n = 199
Camphor
10
0
0
200
400
600
800
1000
1200
(c).
5000
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 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
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).
60
50
40
Control
30
Camphor
20
n = 45
n = 36
10
0
0
10
15
20
25
30
160
35
40
45
50
(c).
5000
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
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
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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
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
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
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.
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
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.
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.
168
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
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.
170
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.
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
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:
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.
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
177
178
References
Abivardi, C. (1979). Fungicidal effect of camphor on 7 soilborne fungi. Phytopathologia
Mediterranea; 18, 213-215.
Adam, P. (2003). Threatened Species Conservation Act Notice of Preliminary
Determination Rejection of a Proposal. NSW Government. Gazette; 186, 10916.
Adam, P. (2004). Camphor Laurel (Cinnanomum camphora) most toxic chemo-types
rejection of key threatening process listing.
http://www.nationalparks.nsw.gov.au/npws.nsf/content/camphor_laurel_ktp; accessed
1/10/04.
Addicott, F.T. (1978). Abscission strategies in the behavior of tropical trees. In: Tropical
Trees as Living Systems. (ed.) P.B. Tomlinson and M.T. Zimmerman, Cambridge
University Press, Cambridge.
Ali, N. A., Mohtar, M., Shaari, K., Rahmanii, M., Ali, A. M. and bin Jantan, I. (2002).
Chemical composition and antimicrobial activities of the essential oils of
Cinnamomum aureofulvum Gamb. Journal of Essential Oil Research; 14, 135-138.
Anon. (1998). Ecology of Weeds: Camphor laurel. Publication of Fletcher Laboratory; Qld.
Available at: http://www.nrm.qld.gov.au/alan_fletcher/ecology/camphor_laurel.html;
accessed 2/3/04.
Ashley, J. and Rushforth, S. R. (1984). Growth of soil algae on top soil and processed oil
shale from the Uintah Basin, Utah, USA. Reclamation Revegetation Research; 3, 4963.
Asplund, R. O. (1968). Monoterpenes: relationship between structure and inhibition of
germination. Phytochemistry; 7, 1995-1997.
Asplund, R. O. (1969). Some quantitative aspects of the phytotoxicity of monoterpenes. Weed
Science; 17, 454-455.
Atsutajingu. (2004). Giant camphor tree. http://www.atsutajingu.or.jp/eng/pre/hozoju.htm;
accessed 15/11/04.
Australian Government Bureau of Meteorology. (2004a) Climate Queensland.
http://www.bom.gov.au/climate/averages/tables/ca_qld_names.shtml; accessed
6/11/04.
179
180
Buscot, F. (2005). What are soils. In: Microorganisms in Soils: Roles in Genesis and
Functions. (eds.) F. Buscot and A. Varma. Springer, Berlin.
Cartwright-Jones, C. (2006). The Henna Page. http://hennapage.com/henna/warnings.html;
accessed 16/3/06.
Chamberlain, C. J. (1925). Methods in Plant Histology. University of Chicago Press,
Chicago, Illinois.
Chao, S., Young, D. G. and Oberg, C. J. (2000). Screening for inhibitory a activity of
essential oils on selected bacteria, fungi and viruses. Journal of Essential Oil
Research; 12, 639-649.
Chapman, A. D. (1991) Australian Plant Name Index. Australian Flora and Fauna Series
Number 12. Australian Biological Resources Study, Canberra, ACTU.
Chemindustry (2006). Chemicals. http://www.chemindustry.com/chemical/142662.html;
accessed 19/3/06.
Cheng, H. (1992). A conceptual framework for assessing allelochemicals in the soil
environment. In: Allelopathy: Basic and Applied Aspects. (eds.) S. J. H. Rizvi and V.
Rizvi, Chapman and Hall, London.
Cheng, C. and Gordon, I. L. (2000). The Richards function and quantitative analysis of
germination and dormancy in meadowfoam (Limnanthes alba). Seed Science
Research; 10, 265-277.
Chen, Z-H., Wu, B., Li, J-Y., Zhao, J-G., Zhou, X-Y. and Zhang, Y-K. (2004). Germination
of the seeds and growth of seedlings of Cinnamomum camphora (L.) Prestl. Plant
Species Biology; 19, 55-58.
Chien, C-T. and Lin, T-P. (1999). Effects of moisture content and temperature on the
storage and germination of Cinnamomum camphora seeds. Seed Science and
Technology; 27,315-320.
Chinese Business World. (2004). Provincial Climate. http://www.cbw.com/general.html;
accessed 6/11/04.
Choi, M. S., Shin, K., Kwon, O. W., Yoon, S. R., Jung, M. Y., Song, K. H., Ahn, J. K., Lee,
W. L., and Son, S. H. (1999). The screening of antimicrobial species from woody
plants. Forest Research Institute. Journal of Forestry Science; 62, 141-154.
Chopra, R. N., Nayar, S. L. and Chopra, I. C. (1956). Glossary of Indian Medicinal Plants.
Council of Scientific & Industrial Research, New Delhi, India.
181
Chou, C-H., Chang, S-J., Cheng, C-M., Wang, Y-C., Hsu, F-H. and Den W-H. (1989). The
selective allelopathic interaction of a pasture-forest intercropping in Taiwan: II.
Interaction between kikuyu grass and three hardwood plants. Plant and Soil; 116, 207215.
Christophel, D. C. and Rowett, A. I. (1996). Flora of Australia Supplementary Series
Number 6. Leaf and Cuticle Atlas of Australian Leafy Lauraceae. Australian Nature
Conservation Agency, Canberra.
Dewey, L. H. (1897). The camphor tree. United States Department of Agriculture, Division of
Botany, Circular No.12, Washington.
Diana, R; Yahata, H. and Nagao, A. (1997). The effects of light and light intensity on the
morphological characteristics of Quercus glauca and Cinnamomum camphora.
Bulletin of the Institute for Tropical Agriculture, Kyushu University; 20, 43-52.
Dunphy, M. (1991) Rainforest weeds of the Big Scrub. In: Rainforest Remnants (ed.)
S. Phillips, NSW National Parks and Wildlife Service, Hurstville.
Edwards, C. A. (2000). Soil invertebrate controls and microbial interactions in nutrient and
organic matter dynamics in natural and agro-ecosystems. In: Invertebrates as
Webmasters in Ecosystems. (eds.). D.C. Colemond, and D.F. Hendrix, Cabi
Publishing, USA.
Firth, D. J. (1979). Ecology of Cinnamomum camphora (L.) Nees & Eberm (Camphor
laurel) in the Richmond-Tweed Region of North-Eastern New South Wales.
Litt.B. Thesis, University of New England, Armidale, NSW.
Firth, D. J. (1981). Camphor laurel (Cinnamomum camphora) a new weed in north-eastern
New South Wales. Australian Weeds; 1, 26-28.
Floridata. (2004). Cinnamomum camphora. http://floridata.com/ref/c/cinn_cam.cfm;
accessed 15/11/04.
Frizzo, C. D., Santos, A. C., Paroul, N., Serafini, L. A., Dellacassa, E., Lorenzo, D. and
Moyna, P. (2000). Essential oils of camphor tree (Cinnamomum camphora Nees &
Eberm) cultivated in southern Brazil. Brazilian Archives of Biology and Technology;
43 (3), 313-316.
Ghosh, R. C. (1977). Handbook of Afforestation Techniques. Controller of Publications,
Delhi, India.
Goldstein, H. (1995). Multilevel Statistical Models (2nd ed). Arnold Publishing.
Grieve, M. (1931). A Modern Herbal. Penguin Books, England.
Grime, J. P. (1979). Plant Strategies and Vegetation Processes. John Wiley & Sons,
Chichester.
Hadington, P. W. and Johnston, J. A. (1979) Australian Trees. A Guide to Their Care and
Cure. (3rd Ed). NSW University Press Ltd, Kensington, Australia.
Harden, G. J. (1990). Lauraceae. In: Flora of New South Wales Vol. 1. (ed) Harden, G.J.
NSW University Press Ltd, Kensington, Australia.
183
Hartmann, H. T. and Kester, D. E. (1968). Plant Propagation: Principles and Practices. (2nd
Ed). Prentice Hall, New Jersey, USA.
Hiroshima City-Business Placement. (2004). Official tree of Hiroshima City camphor tree.
http://business.econ.city.hiroshima.jp/english/photo1.html; accessed 15/11/04.
Hirota, N. and Hiroi, M. (1967). The later studies on the camphor tree, on the leaf oil of each
practical form and its utilization. Journal of Essential Oil Research; 58, 364-367.
Holmes, G. (1987). Avifauna of the Big Scrub. National Parks and Wildlife Service, N.S.W.,
Hurstville.
Howarth, W. O. and Warne L. G. G. (1952). Practical Botany. University Tutorial Press Ltd,
London.
Huadong, R., Xiaohua, Y., Yinxiang, S., Lianzhong, Z. and Jianshi, C. (2000). Study of
biomass variance and comprehensive evaluation at the seedling stage of
Cinnamomum camphora provenances. Forest Research; 13 (1), 80-85.
Huergo, H. H. and Retamar, J. A. (1978). Aceite escencial de Cinnanomum camphora.
Rivista. Ital. Essenze, Profumi, Piante Off; Araomat; Syndets, Saponi, Cosmet.
Aerosol; 60 (11), 637-638.
Immelman, W. F. E., Wicht, C. L. and Acherman, D. P. (1973). Our Green Heritage.
Tafelberg. Cape Town, South Africa.
Issa, M., Defarge, C., Bissonnais, Y., Marin, B., Duval, O., Bruad, A., DAcque, L.,
Nordenberg, S. and Annerman M. (2007). Effects of the inoculation of cyanbacteria
on the microstructure and the structural stability of a tropical soil. Plant and Soil; 290,
(1-2).
Jantan, I. and Goh, S. H. (1992). Essential oils of Cinnamomum species from peninsular.
Malaysia. Journal of Essential Oil Research; 4, 161-171.
Japan Broadcasting Corporation. (2004). Kusunoki (a camphor tree) of Tottoriyamadashrine. http://www.nhk-chubu-brains.co.jp/DDT-E/mie/toin/kusunoki.html; accessed
15/11/04.
Johansen, J. R., Kasper, K., St. Clair, L. L., Warren, S. and Pendleton, B. (1994).
Stabilization of damaged soil crust communities using cyanobacterial amendments:
development of inocula and field testing. American Journal of Botany; 81 (6), 108.
Johns, G. W. (1994). Allelopathic Interactions Between Pennisetum clandestinum and
Cinnamomum camphora and Other Sub-Tropical Tree Species. Project paper
B.App. Science (Cons Tech), Southern Cross University, Lismore, NSW.
184
Kanowski, J., Catterall, C. P., Dennis, A. J. and Westcott, D. A. (Eds.) (2004). Animal-plant
interactions in conservation and restoration, Cooperative Research Centre for
Tropical Rainforest Ecology and Management. Rainforest CRC, Cairns.
Knusel, L. (1998). On the accuracy of statistical distributions in Microsoft Excel 97.
Comparative Statistics and Data Analysis; 26, 375-377.
Kong, J-O., Park, I-K., Choi, K-S., Shin, S-C. and Ahn, Y-J. (2007). Nematicidal and
propagation activities of Thyme red and white oil compounds towards
Bursaphelenchus xylophilus (Nematoda: Parasitaphelenchidae). Journal of
Nematology; 39(3), 237-242.
Labs-Mart. (2006). Reagents. http://www.labs-mart.com/Reagents.htm; accessed 18/3/06.
Laessle, A.M. (1958). A report on succession studies of selected plant communities on the
University of Florida Conservation Reserve, Welaka, Florida. Journal of Florida
Academy of Science; 21, 101-112.
Lange, O. L., Meyer, A., Zellner, H. and Heber, U. (1994). Photosynthesis and water
relations of lichen soil crusts: field measurements in the coastal fog zone of the
Namib Desert. Functional Ecology; 8, 253-264.
Lawrence, B. M. (1995). Progress in essential oils. Perfume and Flavor; 20 (4), 2941.
Le Cussan, J., Hyland B. P. M. and Weber, J. Z. (2007). Lauraceae. In: Flora of Australia
Vol. 2. Winteraceae to Plantanaceae. (ed.) A. Wilson. Australian Biological Resources
Study, Canberra, ACT.
Leicach, S. R., Sampietro, D. A. and Narwal, S. S. (2009). Allelochemicals: Role in PlantEnvironment Interactions. Studium Press, Houston, USA.
Liao, J-C. (1996). Lauraceae. In: Flora of Taiwan (2nd Ed.) Vol. 2. (ed.) T-C Huang, Dept.
Botany, National Taiwan University, Taipei, Taiwan.
Liengme, B. V. (2000). A Guide to Microsoft Excel for Scientists and Engineers.
Butterworth-Heinemann, Oxford.
Liu, C. H; Mishra, A. K. and Tan, R. X. (2001). Composition and antifungal activity of
essential oils from Artemisia princes and Cinnamomum camphora. International Pest
Control; 43 (2), 72-74.
Liu, R., Wei, G., Qing, Y., He, W. and Liu, W. (2002). Cinnamomin, a type II ribosomeinactivating protein, is a storage protein in the seed of the camphor tree
(Cinnamomum camphora). Biochemistry Journal; 362, 659-663.
185
186
Neilan, W. (2004). Seeing the Forest through the Weeds: An Investigation into the Potential
for Rainforest Regeneration in Camphor Laurel Dominated Regrowth in the Big
Scrub. Honours Thesis, Griffith University, Brisbane.
Neish, A. C. (1964). Major pathways of biosynthesis of phenols. In: Biochemistry of
Phenolic Compounds. (ed.) J. B. Marborne, Academic Press, New York: 295-359.
Pakenham, T. (2003). Remarkable Trees of the World. Weidenford and Nicholson, London.
Panetta, F. D. (2001) Seedling emergence and seed longevity of the tree weeds Celtis
sinensis and Cinnamomum camphora. Weed Research; 41, 83-95.
Paul, M., Catterall, C. P., Pollard, P. C. and Kanowski, J. (2010). Does soil variation between
rainforest, pasture and different reforestation pathways affect the early growth of
rainforest pioneer species? Forest Ecology and Management (in press).
Pelissier, Y., Marion, C. and Prunac, S. (1995). Volatile components of leaves, stems and
bark of Cinnamomum camphora Nees et Ebermaier. Journal of Essential Oil
Research; 7, 313-315.
People Daily Online. (2004). 2,000-year-old camphor tree found in east China.
http://english.people.com.cn/200310/23/eng20031023_126712.shtml; accessed
15/11/04.
Philippot, L. and Germon, J. C. (2005). Contribution of bacteria to Initial Input and Cycling
of Nitrogen in Soils. In: Microorganisms in Soils: Roles in Genesis and Functions.
(eds.) F. Buscot and A. Varma. Springer, Berlin.
Pianka, E .R. (1970). On r- and K-selection. American Naturalist; 104, 592-597.
Pino, J. A. and Fuentes, V. (1998). Leaf oil of Cinnamomum camphora (L.) J. Presl. from
Cuba. Journal of Essential Oil Research; 10, 531-532.
Pizzolato, P. (1964). Histochemical recognition of calcium oxalate. J. Histochem. Cytochem;
12: 333-336.
Rahman, A. U., Choudhary, M. I., Faroo, A., Ahmed, A., Iqbal, M. Z., Demirci, B., Demirci,
F. and Husnu Can Baser, K. (2000). Antifungal activities and essential oil
constituents of some spices from Pakistan. Journal of the Chemistry Society of
Pakistan; 22 (1), 60-65.
Ranasinghe, L., Jayawardena, B. and Abeywickrama, K. (2002). Fungicidal activity of
essential oils of Cinnamomum zeylanicum (L.) and Syzygium aromaticum (L.) Merr et
L.M. Perry against crown rot and anthracnose pathogens isolated from banana.
Letters in Applied Microbiology; 35, 208-211.
187
189
Stubbs, B. J. and Brushett, D. (2001). Leaf oil of Cinnamomum camphora (L.) Nees and
Eberm. from Eastern Australia. Journal of Essential Oil Research; 13, 51-54.
Stubbs, B. J., Specht, A. and Brushett, D. (2004). The essential oil of Cinnamomum
camphora (L.) Nees and Eberm. variation in oil composition throughout the tree in
two chemotypes from eastern Australia. Journal of Essential Oil Research; 16, 200205 (May/June).
Swarbrick, J. T. (1991). Towards a rating scheme for environmental weeds. Plant Protection
Quarterly; 6, 185.
Tiwari, R. and Dixit, V. (1994). Fungitoxic activity of vapours of some higher plants
against predominant storage fungi. National Academy of Science Letters India; 17 (34), 55-57.
Torrsell, K. B. G. (1997). Natural Product Chemistry: A mechanistic, biosynthetic and
ecological approach. Swedish Pharmaceutical Press, Stockholm, Sweden.
Troup, R. S. (1921). The Silviculture of Indian Trees. Vol. III. Lauraceae to Coniferae.
Clarendon Press, Oxford.
United States Department of Agriculture. (1897). The camphor tree. (Cinnamomum
camphora Nees & Eberm). Circular No.12. U.S. Department of Agriculture Division of
Botany.
University of Berkeley. (2006). Staining.
http://microscopy.berkeley.edu/Resources/instructions/staining; accessed 17/3/06.
University of Illinois. (2006). Histochemical Tests.
http://boneslab.bio.ntnu.no/histochemicaltests.htm; accessed 20/3/06.
Valder, P. (1999). The Garden Plants of China. Florilegium, Glebe, Australia.
Vander Willigen, C., Sherwin, H. W. and Pammenter, N. W. (2000). Xylem hydraulic
characteristics of subtropical trees from contrasting habitats grown under identical
environmental conditions. New Phytologist; 145, 51-59.
Whittaker, R. H. (1970). The biochemical ecology of higher plants. In: Chemical Ecology.
(eds.) E. Sandheimer and J.B. Simeone, Academic Press, New York. 43-67.
Whittaker, R. H. and Feeny, P. P. (1971). Allelochemicals: Chemical interactions between
species. Science; 171, 757-770.
190
Whitton, B. A. and Potts, M. (2000). The Ecology of Cyanobacteria: Their Diversity in Time
and Space. Kluwer Academic Publishers, Netherlands.
Woodford, R. (1993). Rainforest regeneration on the north coast: Rocky Creek dam. In:
Bushland in Our Cities and Suburbs. Part 2: Making Bush Regeneration Work.
Proceedings of a seminar held at the Institution of engineers, Milsons Point, Sydney,
October 22 and 23.
Xiang, Y., Huadong, R., Kailiang, W., Kejiu, W. and Sen, G. (2001). A preliminary study on
genetic variance of bud germination at the seedling stage of Cinnamomun camphora
(L.) presl. Acta Agriculturae Universitatis Jiangxiensis; 23 (4), 537-543.
Xiang, Y. and Liqing, Z. (2001). Application of geostatistics method to study on growth
differentiation at the seedling stage of Cinnamomun camphora. Acta Agriculturae
Universitatis Jiangxiensis; 23 (3), 350-354.
Xiaohua, Y., Huadong, R., Kejiu, W., Aisheng, W., Sen, G. and Bingsheng, L. (1999). The
study on genetic variance at seedling stage of Cinnamomun camphora L. Acta
Agriculturae Universitatis Jiangxiensis; 21 (3), 320-328.
Yurugi, Y. and Nagata, H. (1981). Studies on dormancy in woody Plants (II). Growth
patterns and distribution of evergreen broad-leaf trees. Bulletin of the Faculty of
Agriculture Mie University; 63, 199-203.
Zhang, D-H., Hong, C-H., Shen, X-Z., Ye, G-F., Zhang, R-F., Shi, M-F., Shi, M-R. and
ZHu, Y-X. (2002). Experimental study on salinity tolerance in tree species in Shoal of
Zhejiang. Journal of Zheijiang Forestry, Science and Technology; 22 (2), 16-21.
Zhou, X., Li, X. D., Yuan, J. Z., Tang. Z. H. and Liu, W-Y. (2000). Toxicity of cinnamomin
a new type II ribosome-inactivating protein to bollworm and mosquito. Insect
Biochemistry and Molecular Biology; 30, 259-264.
Zhu, L., Li, Y., Li, B. and Xia, N. (1993). Aromatic Plants and Essential Oil Constituents.
South China Institute of Botany, Chinese Academy of Sciences, Hai Feng Publishing
Co; Hong Kong.
Zhu, L., Ding, D. and Lawrence, B.M. (1994). The Cinnamomum species in China: resources
for the present and future. Perfume and Flavor; 19 (4), 17-22.
191
APPENDICES
September 2009
LIST OF APPENDICES
A1
A2
A3
A4
A5
A6
A7
A8
A9
A10
A11
A12
A13
A14
A15
ii
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
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
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
% 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
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
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
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
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
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
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
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
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
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
Control
100
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
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
90
80
% germination
70
60
50
40
30
20
10
0
1
Figure.
Figure
A1.33 X Germination of *Cinnamomum camphora from
depulped seed of fallen ripe fruit.
100
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
100
% 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
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
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
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
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
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
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
A2-3
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
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
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
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
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
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
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
A3-5
99
105 112
vines/twinners (species n = 7)
43%
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
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
FigureFigure
A4.2 Propagule
weights
in theinresponse
groups.groups.
weights
the response
X. Propagule
35
30
25
32%
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
35
30
29%
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%
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
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
% 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
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
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
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
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
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
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
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
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
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.
100
observed germination
90
fitted germination
% germination
80
70
60
50
40
30
r2 = 0.937535
20
10
0
1
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
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
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
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
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
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
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
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
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
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
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
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
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
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
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
'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
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
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
Days
100
observed germination
90
fitted germination
% germination
80
70
60
50
40
30
20
r = 0.985642
10
0
1
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
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
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
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
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
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
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
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
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
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 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
'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.
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
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
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
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
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
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
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
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
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
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
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
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.
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
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.
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
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.
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)
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.
Appendix 11.
for shoot length, radicle length, leaf number and leaf area
of each species at completion of the germination triali
Control
Cineole
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
2.22916639
1.66140353
Observations
Pooled Variance
df
0.014105474
t CritiCamphorl one-tail
155.954475
93
t Stat
P(T<=t) one-tail
Pooled Variance
df
Observations
136.7410803
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
0.028210948
0.899452142
0.058750145
*No Sig difference exists in shoot length between Cineole and Camphor
A11-2
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
0.99420737
t Stat
0.161350851
1.66140353
Cineole
32.39130435
Pooled Variance
93
Camphor
Mean
109.9753083
Observations
Pooled Variance
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
0.322701702
0.851974186
0.334502216
*No Sig difference exists in rad. length between Cineole and Camphor
Control
Mean
Control
1.826086957
1.795918367
Mean
1.61352657
1.040816327
Variance
46
49
Variance
Observations
Pooled Variance
Cineole
1.317934186
Cineole
Camphor
1.403846154
Mean
1.795918367
1.403846154
1.61352657
0.951357466
Variance
1.040816327
0.951357466
46
52
49
52
1.261749234
93
Camphor
1.826086957
Observations
Pooled Variance
df
Pooled Variance
df
Observations
0.994731459
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
0.898422255
0.066359238
0.051113313
A11-3
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
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
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
0.347236269
0.114788362
0.437376178
Control
Cineole
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
t Stat
Pooled Variance
df
Observations
35.67764019
66
-3.328490732
t Stat
Pooled Variance
df
Observations
46.08777007
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
0.001431157
0.410187677
7.01284E-05
A11-4
Radicle
Length
Length
Allocasuarina littoralis - Radical
t-Test: Two-Sample Assuming Equal Variances
Control
Cineole
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
66
df
12.30294641
Observations
Pooled Variance
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
0.377828878
0.412850144
-0.031990577
0.974552843
No dif in phyllode no.between Control and Cineole, Control and Camphor, Cineole and Camphor
A11-5
Control
Cineole
Control
Mean
51.30769231
38.6
Variance
126.0641026
239.6
13
10
Observations
Pooled Variance
174.7223443
Cineole
51.30769231
52.66666667
Mean
Variance
126.0641026
363.3333333
Variance
13
12
Observations
239.5406912
21
Camphor
Mean
Pooled Variance
df
df
52.66666667
239.6
363.3333333
10
12
Observations
Pooled Variance
307.6533333
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
0.828321623
t Stat
20
P(T<=t) one-tail
-0.219338015
Camphor
38.6
-1.873008147
0.03787938
0.075758761
A11-6
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
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
0.086766525
Cineole
40.38461538
Pooled Variance
21
Camphor
Mean
133.2497213
Observations
Pooled Variance
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
0.421505401
0.055900074
Control
Cineole
Control
Mean
2.076923077
0.7
Variance
4.076923077
2.455555556
13
10
Observations
Pooled Variance
3.382051282
Cineole
2.076923077
2.416666667
Mean
Variance
4.076923077
3.356060606
Variance
13
12
Observations
3.732162765
21
Camphor
Mean
Pooled Variance
df
df
2.416666667
2.455555556
3.356060606
10
12
Observations
Pooled Variance
2.950833333
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
0.664542378
t Stat
20
t Stat
-0.439302099
Camphor
0.7
-2.333957355
0.03014321
2.085962478
0.03014321
A11-7
Control
Mean
Variance
Control
1.9
26.33333333
11.65555556
13
10
Observations
Pooled Variance
Cineole
20.04285714
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
11.65555556
14.90909091
10
12
Observations
13.445
0
df
t Stat
1.713870006
1
2.068654794
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
Camphor
1.9
0.5
Mean
Pooled Variance
23
P(T<=t) two-tail
2.079614205
26.33333333
df
t Stat
Cineole
5
21
Camphor
5
Observations
Pooled Variance
df
0.062292984
Control
Cineole
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
t Stat
t Stat
0.00183779
1.665150648
0.00367558
1.991675163
0.00367558
49.87863058
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
0.179361912
Observations
Pooled Variance
df
2.997561399
P(T<=t) two-tail
t CritiCamphorl two-tail
23.98825011
76
P(T<=t) one-tail
t CritiCamphorl one-tail
Pooled Variance
df
Observations
3.82406E-05
A11-8
Radicle
Length
Length
Alpinea caerulea - Radical
t-Test: Two-Sample Assuming Equal Variances
Control
Cineole
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
Pooled Variance
77
df
5.531649761
2.0973E-07
1.664884621
4.1946E-07
1.991256795
4.1946E-07
Observations
176.3471061
Observations
Pooled Variance
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
0.693736363
-7.823303072
1.0621E-11
1.987291398
1.0621E-11
Control
Mean
Variance
Control
1.94
0.892857143
Mean
0.465714286
1.876984127
Variance
50
28
Observations
Pooled Variance
Cineole
0.967086466
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
76
t Stat
Camphor
1.94
Observations
Pooled Variance
df
0.727835672
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
2.30873E-05
0.098015306
-4.454071136
2.46575E-05
A11-9
Control
Cineole
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
77
df
Observations
Pooled Variance
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
t Stat
0
88
t Stat
-2.831501093
5114.475545
0.005502249
-5.1864817
1.35641E-06
Control
Cineole
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
Pooled Variance
df
Observations
59.34312494
128
17.94915254
20.1
Variance
67.32495617
48.09130435
59
70
Observations
Pooled Variance
df
56.87517683
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
0.004116366
0.103683128
-1.613720816
0.109070009
*No Sig difference exists in shoot length between Cineole and Camphor
A11-10
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
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
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
0.424278795
0.047090415
0.253390731
*No Sig difference exists in rad. length between Cineole and Camphor
Control
Cineole
Control
Mean
1.591549296
1.55
Variance
0.245070423
0.251694915
71
60
Observations
Pooled Variance
0.248100229
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
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
0.226339455
A11-11
0.250310559
60
df
1.773856757
1.442857143
0.251694915
139
Camphor
1.55
Observations
Pooled Variance
df
Cineole
1.591549296
129
Camphor
Mean
Pooled Variance
df
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
129
df
5709.273185
Observations
Pooled Variance
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
0.332704115
*No Sig difference exists in leaf area between Control and Cineole
0.27321021
1.977305146
0.27321021
0.892683683
Control
Mean
Variance
Cineole
Pooled Variance
44.03333333
Mean
57.30555556
36.99885057
Variance
28
30
46.78958333
-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
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
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
0.839639259
0.56621587
2.002466317
0.56621587
A11-12
Camphor
Mean
df
P(T<=t) one-tail
Cineole
42.75862069
56
Camphor
43.25
Observations
Pooled Variance
df
t Stat
Control
43.25
Observations
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
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
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
0.215150852
0.176398074
0.017704611
Control
Cineole
Control
Mean
15.60714286
11.3
Variance
191.8029101
91.18275862
28
30
Observations
Pooled Variance
139.6960459
Cineole
15.60714286
15.4137931
Variance
191.8029101
135.7512315
28
29
Observations
163.2675101
56
Camphor
Mean
Pooled Variance
df
Variance
15.4137931
91.18275862
135.7512315
30
29
113.0760436
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
0.170989829
0.954662237
0.142905175
A11-13
-1.485561939
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
55
df
1.37330344
t Stat
11.94120429
Observations
Pooled Variance
9.118047994
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
0.175230748
0.337664744
0.013819183
Camphor
Mean
12.90769231
14.59649123
Variance
18.77259615
14.60213033
65
57
Observations
Pooled Variance
16.82637877
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
0.025067145
A11-14
Radicle
Length
Length
Callistemon viminalis - Radical
Control
Camphor
Mean
13.06153846
12.66666667
Variance
52.96490385
64.44047619
65
57
Observations
Pooled Variance
58.32017094
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
0.776177531
Control
Mean
Variance
1.719298246
0.860863095
0.884085213
64
57
Observations
Pooled Variance
0.871791151
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
4.80939E-27
A11-15
Control
Camphor
Mean
3.661290323
1.456140351
Variance
7.604706504
1.502506266
62
57
Observations
Pooled Variance
4.683995279
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
1.78689E-07
Control
Cineole
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
t Stat
Pooled Variance
df
Observations
501.9885147
111
-0.522537831
t Stat
Pooled Variance
df
Observations
530.3613625
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
0.602337283
2.92834E-05
1.98044745
0.000631268
A11-16
Observations
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
101
df
-2.774080434
t Stat
339.8999904
Observations
Pooled Variance
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
0.006595173
2.38968E-06
0.401108391
*No Sig difference exists in rad. length between Cineole and Camphor
Control
Cineole
Control
Mean
2.225806452
2.254901961
Variance
0.374405077
0.55372549
62
51
Observations
Pooled Variance
0.455180038
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
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
117
1.98044745
0.378643635
A11-17
0.415057068
df
-1.284118322
2.367647059
0.55372549
128
Camphor
2.254901961
Observations
Pooled Variance
df
-0.228126674
Cineole
2.225806452
111
Camphor
Mean
Pooled Variance
df
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
111
df
65056.64467
250.3921569
194.761194
Variance
42235.64314
24788.21483
51
67
Observations
Pooled Variance
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
0.184947086
0.005709101
1.665476857
0.098516164
Control
Cineole
Control
Mean
57.24137931
55.80645161
Variance
274.2610837
506.827957
29
31
Observations
Pooled Variance
394.554294
Cineole
57.24137931
70.74074074
Mean
Variance
274.2610837
253.2763533
Variance
29
27
Observations
264.1573246
58
Camphor
Mean
Pooled Variance
df
df
70.74074074
506.827957
253.2763533
31
27
Observations
Pooled Variance
389.1075696
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
0.003021927
t Stat
56
t Stat
-3.105766235
Camphor
55.80645161
-2.876063348
0.005685912
A11-18
Radicle
Length
*Cassia coleutioides - Radical
Length
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
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
0.266801183
0.428248755
-0.311586023
0.756512235
Control
Cineole
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
Pooled Variance
df
Observations
0.256279268
58
Pooled Variance
df
Observations
0.260752688
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
0.029828972
0.431591625
-1.386543424
0.171076448
A11-19
Control
Mean
Variance
Cineole
63.58064516
Mean
2474.455172
2171.051613
Variance
30
31
Observations
Pooled Variance
Control
94.6
2320.182176
Camphor
Mean
63.58064516
62.55555556
2474.455172
1264.717949
Variance
2171.051613
1264.717949
30
27
31
27
1902.579394
59
Cineole
62.55555556
Observations
Pooled Variance
df
Camphor
94.6
Pooled Variance
df
Observations
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
0.014667307
0.007641512
0.926171597
Cineole
Camphor
Mean
143.3870968
97.7173913
Variance
1210.957223
746.3807931
93
92
Observations
Pooled Variance
979.9383426
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
0.452714098
A11-20
Radicle
Length
Length
*Cinnamomum camphora (fresh from tree) - Radical
0 value for Control - stats not possible
Cineole
Camphor
Mean
66.79569892
52.55434783
477.2730248
168.4036073
93
92
Variance
Observations
Pooled Variance
323.6822215
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
2.22317E-07
Cineole
Camphor
Mean
3.935483871
3.77173913
1.082748948
0.661610129
93
92
Variance
Observations
Pooled Variance
0.873330191
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
0.234964268
A11-21
Cineole
Camphor
Mean
478.9784946
372.7826087
71987.69518
44700.28189
93
92
Variance
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
Control
Cineole
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
df
t Stat
Observations
Pooled Variance
3283.205128
21
-1.222003134
209
172.5
Variance
610
2491.666667
10
Observations
Pooled Variance
df
1080.416667
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
0.235246464
0.716701383
0.085037025
A11-22
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
93.75
Variance
3414.423077
256.25
13
Observations
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
0.018405971
0.70510843
2.131450856
0.70510843
-3.448408431
0.004819113
Control
Cineole
Control
Mean
10.92307692
12.9
Variance
21.24358974
25.87777778
13
10
Observations
Pooled Variance
23.22967033
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
15
df
-0.228721262
Camphor
12.9
Observations
Pooled Variance
df
-0.975160286
Cineole
10.92307692
21
Camphor
Mean
Pooled Variance
df
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
0.340580945
0.626814723
0.822174575
value > 0.05 - null hypothesis (Ho) accepted
A11-23
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
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
0.258972205
0.986393458
-0.776102531
0.452714098
Control
Cineole
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
Pooled Variance
df
141.0968912
134
t Stat
Observations
2.38619454
35.3375
Variance
197.7509615
128.277057
65
80
Observations
159.370273
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
0.018421902
4.82177249
Camphor
Mean
3.47765E-06
1.65558049
0.065779895
*No Sig difference exists in shoot length between Cineole and Camphor
A11-24
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
134
df
243.1801163
31.69230769
29.525
Variance
224.1538462
153.6196203
65
80
Observations
Pooled Variance
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
0.112453873
0.008201408
0.95374559
0.170910923
1.65558049
0.341821845
Control
Cineole
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
Pooled Variance
df
Observations
0.229633205
133
0.738461538
0.75
Variance
0.321153846
0.240506329
65
80
Observations
Pooled Variance
df
0.276600323
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
0.006282039
0.003115301
-0.131383019
0.447828492
1.65558049
0.895656984
A11-25
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
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
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
t Stat
18558.72782
P(T<=t) one-tail
4.34168034
Camphor
Mean
2.60866E-05
-0.318083772
0.750882217
Control
Mean
Variance
Control
26.5
24.03571429
Mean
16.92028986
28.51678141
Variance
70
84
Observations
Pooled Variance
Cineole
23.25258459
Cineole
Camphor
22.24390244
Mean
24.03571429
22.24390244
16.92028986
13.73902439
Variance
28.51678141
13.73902439
70
41
84
41
15.75285299
152
Camphor
26.5
Observations
Pooled Variance
df
Pooled Variance
df
Observations
23.71100677
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
0.001917729
3.12252E-07
0.055721963
*No Sig difference exists in shoot length between Cineole and Camphor
A11-26
Control
Mean
Variance
29.85714286
Mean
118.5507246
140.9913941
Variance
70
84
130.8045113
Observations
Pooled Variance
df
t Stat
Control
24
Observations
Pooled Variance
Cineole
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
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
0.001876875
0.834699385
0.018679597
Control
Cineole
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
df
Observations
Pooled Variance
0.149844964
152
0.87804878
Variance
0.109756098
84
41
Observations
Pooled Variance
df
0.03569304
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
0.000225597
0.000196912
0.000945564
A11-27
Control
Cineole
Control
Mean
4.385714286
1.535714286
Variance
5.660662526
2.90232358
70
84
Observations
Pooled Variance
4.154464286
1.536585366
Mean
Variance
5.660662526
1.954878049
Variance
70
41
Observations
4.300741617
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
7.22671E-15
2.35125E-10
-0.002838782
0.997739582
Control
Cineole
Control
Mean
65.38383838
58.49484536
Variance
184.4430014
160.044244
99
97
Observations
Pooled Variance
172.3693895
Cineole
65.38383838
69.40625
Variance
184.4430014
293.8648026
99
96
Observations
238.3034735
194
Camphor
Mean
Pooled Variance
df
Variance
69.40625
160.044244
293.8648026
97
96
226.6042077
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
0.070445497
t Stat
191
t Stat
-1.819102757
Camphor
58.49484536
Observations
Pooled Variance
df
Mean
-5.034884424
1.10172E-06
A11-28
Control
Cineole
Control
Mean
54.54545455
47.2371134
Variance
197.4953618
79.93277491
99
97
Observations
Pooled Variance
Hypothesized Mean difference
df
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
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
2.34904E-05
1.97233021
1.60068E-07
1.48813704
0.138363898
Control
Cineole
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
df
Observations
Pooled Variance
0.368632896
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
191
-1.059607128
0.14533075
1.652870196
0.2906615
1.972462087
0.2906615
A11-29
t Stat
0.46677827
df
t Stat
0.299193407
193
Camphor
Mean
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
Observations
Pooled Variance
194
df
3659.320956
Observations
Pooled Variance
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
0.062021015
1.97233021
*No sig dif exists in leaf area between Control and Cineole
0.100042735
*No sig dif exists in leaf area between Control and Camphor
-0.297467129
0.766433187
*No sig dif exists in leaf area between Cineole and Camphor
Control
Cineole
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
t Stat
Pooled Variance
df
Observations
73.40128558
21
-0.235676559
Pooled Variance
df
Observations
69.24264706
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
0.815966866
2.03451691
0.007277546
0.001380161
A11-30
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
107.3705234
Mean
Variance
Observations
Pooled Variance
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
-0.447957071
Cineole
21.54545455
Pooled Variance
21
Camphor
Mean
0.658771376
2.03451691
0.046734707
0.016060817
Control
Cineole
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
Pooled Variance
df
Observations
0.541781451
21
Pooled Variance
df
Observations
0.948529412
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
0.291347758
0.00515567
0.472783902
A11-31
Control
Cineole
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
21
df
-0.235676559
18.3503214
Observations
Pooled Variance
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
0.815966866
2.03451691
0.007277546
0.001380161
Control
Cineole
Control
Mean
75.25806452
57.5
Variance
142.1500701
154.2977528
93
90
Observations
Pooled Variance
148.1232401
t Stat
1.653315849
P(T<=t) two-tail
1.19488E-18
t CritiCamphorl two-tail
1.97315785
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
184
df
-0.291261937
t Stat
Camphor
57.5
Observations
Pooled Variance
df
9.867819843
Cineole
75.25806452
181
P(T<=t) one-tail
Camphor
Mean
Pooled Variance
df
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
0.771179069
1.97315785
6.94128E-21
* no significant difference exists in shoot length between the Control 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.
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
t Stat
Pooled Variance
df
Observations
199.7577139
181
df
-0.180262271
Observations
Pooled Variance
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
0.857148338
0.554964386
1.97315785
0.437223777
Control
Mean
Control
4.139784946
3.233333333
Mean
0.77372604
1.282022472
Variance
93
90
Variance
Observations
Pooled Variance
Cineole
1.023661855
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
184
t Stat
Observations
Pooled Variance
df
6.05903624
Cineole
4.075268817
181
Camphor
4.139784946
Observations
Pooled Variance
df
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
7.75535E-09
0.645424124
1.97315785
3.69074E-07
* 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
A11-33
* A significant difference exists in leaf No. exists between the Control and Cineole treatments.
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
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
7.86033E-10
1.97315785
2.22903E-09
t Stat
Camphor
79.5
t Stat
t CritiCamphorl two-tail
6.48692709
Mean
-0.015428398
1.97315785
0.987707401
* 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
21.63636364
18.265625
Variance
48.88111888
25.84895833
66
64
Observations
Pooled Variance
37.54497736
128
Cineole
19.58441558
Mean
Variance
48.88111888
32.35133288
Variance
66
77
39.97144699
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
0.055014415
32.35133288
64
77
29.40421348
0
df
t CritiCamphorl one-tail
19.58441558
25.84895833
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
Cineole
139
-1.437794598
0.07636992
1.655889719
0.15273984
1.977177817
0.15273984
* 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.
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
-1.270949914
Cineole
20.34848485
Pooled Variance
128
Camphor
Mean
t Stat
56.4188849
Observations
Pooled Variance
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
0.206051393
*No significant effect on rad. length can be attributed to the Cineole treatment.
0.01612052
0.032241039
* Radical length was significantly decreased in the Camphor treatment compared to the Control
A11-34
Control
0.002032263
* Sig difference radical length between Cineole and Camphor treatments - Camphor sig. reduced.
Control
Cineole
Control
Mean
4.590909091
3.96875
Variance
2.245454545
2.094246032
66
64
Observations
Pooled Variance
2.171031605
Cineole
4.590909091
2.831168831
Mean
Variance
2.245454545
1.615857826
Variance
66
77
Observations
1.906097449
128
Camphor
Mean
Pooled Variance
df
df
2.831168831
2.094246032
1.615857826
64
77
Observations
Pooled Variance
1.832681258
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
0.017516152
3.79503E-12
1.95694E-06
*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.
A11-35
* A significant difference in leaf No. exists between the Control and Cineole treatments.
Control
Cineole
Control
Mean
12.96923077
8.671875
Variance
152.8427885
57.36681548
65
64
Observations
Pooled Variance
105.4806916
t Stat
4.077922078
Mean
Variance
152.8427885
12.78332194
Variance
65
77
Observations
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
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
0.018990634
1.43094E-08
5.48833E-06
* 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 leaf Area is significantly reduced in the Camphor treatment indicating effect.
Control
Cineole
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
Pooled Variance
df
Observations
12.82724014
105
13.20652174
10.6
Variance
14.71512183
14.42025316
92
80
Observations
Pooled Variance
df
14.57809463
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
-4.432469453
Camphor
Mean
1.27282E-05
1.66140353
2.54564E-05
4.465659983
P(T<=t) one-tail
7.25191E-06
t CritiCamphorl one-tail
1.653866093
1.45038E-05
A11-36
Control
Cineole
Control
Mean
2.466666667
7.293478261
Variance
0.980952381
16.7810559
15
92
Observations
Pooled Variance
14.67437543
t Stat
5.35
Variance
0.980952381
6.660759494
15
80
Observations
5.805734767
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
1.59615E-05
Mean
Variance
5.35
16.7810559
6.660759494
92
80
12.07809463
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
5.01896E-05
0.000338652
Control
Cineole
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
Pooled Variance
df
Observations
0.248611111
102
0.943820225
0.4125
Variance
0.417262513
0.245411392
89
80
Observations
Pooled Variance
df
0.335967671
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
0.283496956
1.66140353
1.52441E-06
7.6611E-09
1.654029802
1.53222E-08
A11-37
t CritiCamphorl one-tail
5.949825758
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
104
8.164740143
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
0.10535293
4.26647E-06
6.578258977
5.71495E-10
Control
Cineole
Control
Mean
67.56060606
59
Variance
59.63473193
86.69333333
66
76
Observations
Pooled Variance
74.13041126
Cineole
67.56060606
66.94186047
Mean
Variance
59.63473193
101.4201094
Variance
66
86
Observations
83.31311252
140
Camphor
Mean
Pooled Variance
df
df
66.94186047
86.69333333
101.4201094
76
86
Observations
Pooled Variance
94.51693314
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
2.49211E-08
0.679288143
1.97490408
6.33509E-07
A11-38
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
t Stat
Pooled Variance
df
Observations
141
df
2.05576558
t Stat
91.76603242
Observations
Pooled Variance
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
0.041649987
0.435322187
0.003199012
* sg. dif in rad length exist between Cineole and Camphor treatments
*Cineole has reduced mean rad. length
Control
Cineole
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
Pooled Variance
df
Observations
66
0.083788329
141
87
Pooled Variance
df
Observations
0.072220103
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
0.47484407
0.065167704
0.09573373
A11-39
0.09573373
1.974717634
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
141
df
66
86
501.8329222
Observations
Pooled Variance
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
0.00021017
5.17237E-10
0.002541478
Control
Cineole
Mean
Variance
Pooled Variance
7.333333333
Mean
9.333333333
5.066666667
Variance
7.393939394
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
11
Camphor
9
Observations
Pooled Variance
df
t Stat
Control
Observations
5.472096531
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
0.294113719
0.150376373
0.878287341
*No Sig difference exists in shoot length between Cineole and Camphor
A11-40
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
11
df
139.8424908
Observations
Pooled Variance
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
0.351441604
0.038567046
0.405938272
*No Sig difference exists in rad. length between Cineole and Camphor
Control
Mean
Variance
1.333333333
0.8
1.090909091
Mean
Variance
Mean
1.333333333
1.397435897
Variance
13
1.376068376
t Stat
Observations
Pooled Variance
df
-1.720912102
Cineole
4.307692308
11
Camphor
4
Observations
Pooled Variance
df
t Stat
Control
Observations
Pooled Variance
Cineole
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
0.113234042
-0.559502885
4.307692308
0.8
1.221719457
18
Camphor
5
0.582714395
0.221518624
A11-41
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
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
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
0.058591164
2.65749E-05
1.615487428
0.06230525
0.124610501
Control
Cineole
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
Pooled Variance
df
Observations
494.5465141
36
Pooled Variance
df
Observations
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
0.001451658
2.0153675
t CritiCamphorl two-tail
0.273116101
df
t Stat
t CritiCamphorl two-tail
180.4344589
56
-4.532505072
1.5553E-05
1.672522103
3.1106E-05
2.003239388
3.1106E-05
* Sig dif exist in shoot length exists between Cineole and Camphor
* mean shoot length reduced in Cineole below Camphor
A11-42
Radicle
Length
Length
Hymenosporum flavum - Radical
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
568.5213675
0
44.65116279
Mean
Variance
440.0641026
277.8516058
Variance
13
43
Observations
-1.139724983
t Stat
0.130963772
Cineole
42.30769231
Pooled Variance
36
Camphor
Mean
P(T<=t) one-tail
313.8988273
Observations
Pooled Variance
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
0.261927544
0.677674458
0.175432949
Control
Cineole
Control
Mean
3.692307692
Variance
2.564102564
1.416666667
13
25
Observations
Pooled Variance
1.799145299
Cineole
3.692307692
2.837209302
Mean
Variance
2.564102564
1.282392027
Variance
13
43
Observations
1.56721659
36
Camphor
Mean
Pooled Variance
df
2.837209302
1.416666667
1.282392027
25
43
Observations
Pooled Variance
df
Camphor
2
1.331219168
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
0.000737482
0.035388014
-2.885087836
0.002640493
0.005280985
A11-43
Control
Cineole
Control
Mean
98.53846154
25.8
Variance
7572.102564
573.6666667
13
25
Observations
Pooled Variance
2906.478632
91.09302326
Mean
Variance
7572.102564
13537.46733
Variance
13
43
Observations
36
Cineole
98.53846154
Pooled Variance
df
Camphor
Mean
df
12211.83072
Observations
Pooled Variance
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
0.00035309
0.832230963
* Not
exists
between
Control
andand
Camphor
*Nosig
sigdif
difininleaf
leafarea
area
exists
between
Control
Camphor
0.007398056
Control
Cineole
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
t Stat
Pooled Variance
df
Observations
25.75182629
175
-1.146487122
t Stat
Pooled Variance
df
Observations
23.90511662
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
0.253158838
0.000193137
1.97323061
0.006621236
* 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.
A11-44
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
175
df
231.6212795
55.75862069
63.56842105
Variance
343.8131516
220.056439
87
95
Observations
Pooled Variance
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
0.136190858
-3.149777687
0.00191355
0.078777906
value < 0.05 - null hypothesis (Ho) rejected
*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.
Control
Cineole
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
Pooled Variance
df
Observations
0.137016423
143
Pooled Variance
df
Observations
0.177392212
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
0.296590865
0.561060113
0.638619399
* 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 similar in Control and Camphor - no effect on leaf No. due Camphor treatment
A11-45
* No significant difference in leaf No. exists between the Control and Cineole treatments.
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
165
df
946.6869492
Pooled Variance
t Stat
1.246587729
t Stat
0.107157753
P(T<=t) one-tail
0.015270427
1026.332574
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
0.214315506
0.030540854
0.477949483
* 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.
Control
Mean
Variance
Cineole
Pooled Variance
Control
38.59375
26.04166667
Mean
117.7384868
214.8492908
Variance
96
48
Observations
149.8807952
Camphor
31.96875
117.7384868
169.5674342
96
96
143.6529605
142
Cineole
38.59375
Observations
Pooled Variance
df
26.04166667
31.96875
Variance
214.8492908
169.5674342
48
96
Observations
Pooled Variance
df
Camphor
Mean
184.555091
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
4.13512E-08
0.00017425
1.972530299
0.00017425
-2.468044623
0.014773091
A11-46
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
Pooled Variance
df
Observations
138.2832353
142
38.8317757
Variance
207.2695035
134.7072827
48
107
Observations
156.9976381
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
2.49606E-11
2.11697E-08
-3.100299443
0.002301877
Control
Cineole
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
Pooled Variance
df
Observations
1.203207931
142
Pooled Variance
df
Observations
1.174098239
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
1.5009E-09
0.000182869
-3.385326359
0.000458985
1.65558049
0.000917969
A11-47
Control
Cineole
Control
Mean
90.13541667
40.125
Variance
3121.402522
2633.81383
96
48
Observations
Pooled Variance
2960.017532
80.50515464
Mean
Variance
3121.402522
5403.273411
Variance
96
97
Observations
4268.31145
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
6.82302E-07
0.30718273
1.972462087
0.30718273
5403.273411
48
97
df
t Stat
80.50515464
2633.81383
4493.031451
191
Camphor
40.125
143
-3.413663694
0.000417053
1.65558049
P(T<=t) two-tail
0.000834106
t CritiCamphorl two-tail
1.976691237
0.000834106
Control
Cineole
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
Pooled Variance
df
Observations
1073.981558
148
Pooled Variance
df
Observations
904.336592
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
0.002635671
5.0947E-07
1.976932253
5.0947E-07
0.005399411
A11-48
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
1.485215963
0.06980676
Cineole
66.69354839
Pooled Variance
148
Camphor
Mean
410.8526751
Observations
Pooled Variance
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
0.139613519
0.003571775
0.105458719
*No Sig difference exists in rad. length between Cineole and Camphor
Control
Cineole
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
t Stat
Pooled Variance
df
0.689775158
148
0.00016185
Observations
Pooled Variance
df
3.681964175
P(T<=t) one-tail
Observations
0.538241738
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
0.000323701
1.12356E-09
6.92188E-05
A11-49
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
148
df
3.237862621
0.00074331
1.655214419
0.00148662
1.976122803
0.00148662
1159.027806
Observations
Pooled Variance
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
2.39456E-07
0.004126361
Control
Cineole
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
Pooled Variance
df
Observations
9.005952381
61
10.10714286
8.4
Variance
5.876984127
8.685714286
28
15
Observations
Pooled Variance
df
6.836062718
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
7.82269E-06
2.01063358
t CritiCamphorl two-tail
1.19827E-06
2.040597092
0.02388279
2.01954208
0.047765579
A11-50
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
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
61
Cineole
26.4
Observations
Pooled Variance
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
0.000572823
0.00144241
2.01954208
0.417138729
Control
Cineole
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
Pooled Variance
df
Observations
0.457142857
61
1.357142857
1.2
Variance
0.904761905
1.028571429
28
15
Observations
Pooled Variance
df
0.947038328
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
0.005485413
2.01063358
t CritiCamphorl two-tail
0.001901844
0.50466279
0.30824873
2.01954208
0.616497461
A11-51
Control
Cineole
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
Pooled Variance
df
Observations
1.720238095
61
0.678571429
0.6
Variance
0.226190476
0.257142857
28
15
Observations
Pooled Variance
df
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
5.89206E-05
2.01063358
t CritiCamphorl two-tail
0.001413233
0.50466279
0.30824873
2.01954208
0.616497461
Control
Cineole
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
Pooled Variance
df
Observations
342.2019824
114
Pooled Variance
df
Observations
186.829753
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
2.73302E-05
0.528513526
2.75983E-05
A11-52
Radicle
Length
Length
Macaranga tanarius - Radical
t-Test: Two-Sample Assuming Equal Variances
Control
Cineole
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
114
df
-0.190817882
t Stat
Observations
Pooled Variance
263.7956797
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
0.849007689
0.669532097
0.730218403
Control
Cineole
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
44
Pooled Variance
df
Observations
114
63
0.25277606
Pooled Variance
df
Observations
0.23493257
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
0.027298137
0.185410662
0.812801998
*No sig dif in leaf number exists between Control and Camphor
*No sig dif in leaf number exists between Cineole and Camphor
A11-53
Control
Mean
Variance
Cineole
72.92957746
3694.256757
17338.0664
38
71
Observations
Pooled Variance
Control
27.5
12620.11353
Variance
Cineole
3694.256757 15349.03738
38
63
10993.21028
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
15349.03738
71
63
16403.8255
0
df
t Stat
66.79365079
17338.0664
99
0.071081312
132
0.276792705
0.39118612
1.656478616
0.78237224
1.978096407
Camphor
72.92957746
Observations
Pooled Variance
df
Mean
0.78237224
Control
Mean
Variance
Cineole
Pooled Variance
Control
77.6835443
70.90789474
Mean
363.1165206
193.4714035
Variance
79
76
Observations
279.9571495
Cineole
Camphor
68.71794872
Mean
70.90789474
68.71794872
363.1165206
229.5038295
Variance
193.4714035
229.5038295
79
78
76
78
296.7411837
153
Camphor
77.6835443
Observations
Pooled Variance
df
Pooled Variance
df
Observations
211.7246719
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
0.012749288
0.001366847
0.351901867
A11-54
Radicle
Length
Omalanthus populifolius - Radical
Length
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
Control
37.73033708
Observations
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
162
Cineole
37.17948718
Observations
Pooled Variance
Camphor
37.73033708
97.07386653
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
0.036359943
0.35701346
151
0.714026919
0.014182013
Control
Cineole
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
t Stat
Pooled Variance
df
0.285048285
151
0.03969825
Observations
Pooled Variance
df
1.766120093
P(T<=t) one-tail
Observations
0.419765665
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
0.079396499
1.65480742
0.232102814
0.557865449
A11-55
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
152
df
Observations
Pooled Variance
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
0.057364837
0.000695609
0.163314135
Control
Mean
Cineole
Control
28.93506494
23.38043478
28.3773069
34.3262064
77
92
Variance
Observations
Pooled Variance
31.6189228
Mean
Cineole
24.41836735
Mean
28.3773069
24.67883442
Variance
77
98
Variance
Observations
26.3035969
167
Camphor
28.93506494
Pooled Variance
df
df
24.41836735
34.3262064
24.67883442
92
98
Observations
Pooled Variance
29.34857299
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
1.54351E-09
3.35963E-08
0.188509009
*No Sig difference exists in shoot length between Cineole and Camphor
A11-56
Radicle
Length
Oplismenus aemulus - Radical
Length
t-Test: Two-Sample Assuming Equal Variances
Control
Cineole
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
Observations
Pooled Variance
167
df
1040.525164
54.07608696
68.06122449
Variance
502.1589823
1485.89312
92
98
Observations
Pooled Variance
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
t Stat
1009.723936
P(T<=t) one-tail
-2.116898007
Camphor
Mean
0.035698274
-3.031765223
0.002774346
Control
Cineole
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
df
Observations
Pooled Variance
0.285108368
167
Pooled Variance
df
Observations
0.245910344
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
0.00010347
0.05001831
1.973771759
0.05001831
P(T<=t) two-tail
0.009045035
t CritiCamphorl two-tail
1.972662176
0.009045035
A11-57
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
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
0.002975447
Camphor
Cineole
19.79220779
14.31632653
Mean
96.6404648
36.96076162
Variance
77
98
63.17843469
Observations
Pooled Variance
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
1.12456E-05
0.068932327
Control
Mean
Variance
Control
34.5
31.78947368
Mean
41.11764706
41.09774436
Variance
52
57
Observations
Pooled Variance
Cineole
41.10723069
Cineole
Camphor
28.71428571
Mean
31.78947368
28.71428571
41.11764706
13.57142857
Variance
41.09774436
13.57142857
52
57
38.21804511
107
Camphor
34.5
Observations
Pooled Variance
df
Pooled Variance
df
Observations
38.43390735
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
0.029626538
0.023679238
1.99896931
0.220183949
*No Sig difference exists in shoot length between Cineole and Camphor
A11-58
Length
*Paspalum dilatatum - Radical
Radicle
Length
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
148.7851098
0
39.25
Variance
126.4328808
591.6428571
52
Observations
182.5789125
Mean
Variance
Observations
Pooled Variance
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
-0.382220336
Cineole
39.80769231
Pooled Variance
107
Camphor
Mean
0.703056044
0.913833254
0.261580512
0.39724909
0.794498179
*No Sig difference exists in rad. length between Cineole and Camphor
Control
Cineole
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
t Stat
Pooled Variance
df
Observations
0.264700212
107
4.721675459
Pooled Variance
df
Observations
0.168324036
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
7.12981E-06
1.17359E-06
4.255540231
1.99896931
7.17799E-05
A11-59
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
107
df
0.8330873
330.996337
Observations
Pooled Variance
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
0.406651228
0.263958116
-1.606938716
1.99896931
0.113149245
Control
Mean
Variance
Control
65.75
41.11494253
Mean
136.8762626
101.0098904
Variance
100
87
Observations
Pooled Variance
Cineole
120.2032463
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
185
Camphor
65.75
Observations
Pooled Variance
df
80.04378928
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
9.23057E-35
1.97315785
1.98417E-31
-2.426410306
0.016306608
A11-60
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
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
1.97287136
t CritiCamphorl two-tail
0.051010791
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
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
8.43224E-06
Control
Mean
Variance
Control
2.23
1.988505747
Mean
0.199090909
0.104517509
Variance
100
87
Observations
Pooled Variance
Cineole
0.155127058
Cineole
Camphor
1.987951807
Mean
1.988505747
1.987951807
0.199090909
0.012048193
Variance
0.104517509
0.012048193
100
83
87
83
0.114353325
185
Camphor
2.23
Observations
Pooled Variance
df
Pooled Variance
df
Observations
0.059383676
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
4.45338E-05
1.97315785
3.02238E-06
0.988197306
A11-61
Control
Cineole
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
168
df
53.68654551
Observations
Pooled Variance
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
0.013865234
t Stat
0
158
t Stat
t CritiCamphorl two-tail
-0.240751288
79.85921683
0.8100681
1.975486157
0.8100681
-2.592271654
P(T<=t) two-tail
0.010428593
t CritiCamphorl two-tail
1.975090527
0.010428593
Control
Cineole
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
Pooled Variance
df
Observations
298.7614849
144
Pooled Variance
df
Observations
208.3832794
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
0.000481233
1.61176E-09
0.006225472
A11-62
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
154
df
-0.890121613
t Stat
0.187394512
1.65480742
169.9391649
Observations
Pooled Variance
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
0.374789025
0.358991289
0.992239083
*No Sig difference exists in rad. length between Cineole and Camphor
Control
Mean
Variance
4.276923077
Mean
4.222222222
5.953365385
Variance
91
65
4.941658342
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
151
t Stat
0.222105638
Camphor
Mean
df
-0.767075066
Cineole
3.596774194
154
Camphor
4
Observations
Pooled Variance
df
P(T<=t) one-tail
Control
Observations
Pooled Variance
Cineole
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
0.444211276
0.215385963
0.079225425
A11-63
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
154
df
180.8289907
Observations
Pooled Variance
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
0.220786711
0.125079513
0.688084886
Control
Cineole
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
Pooled Variance
df
19.92390597
168
t Stat
Observations
2.21691054
t Stat
Pooled Variance
df
Observations
31.56221608
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
0.027971616
0.005433804
7.87504E-05
A11-64
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
df
Camphor
Mean
35.06329114
40.94594595
157.3826618
189.5038874
Variance
237.0856865
189.5038874
91
74
79
74
171.7682414
168
Cineole
40.94594595
Observations
Pooled Variance
Camphor
37.8021978
Pooled Variance
df
Observations
214.0825651
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
0.203196855
-1.532395596
t Stat
151
0.127363416
-2.485228956
0.01403731
1.975799933
0.01403731
Control
Cineole
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
Pooled Variance
df
0.09190873
169
t Stat
Observations
3.3636203
Pooled Variance
df
Observations
0.088089605
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
0.000951754
0.00045393
1.974808583
0.00045393
0.860287098
A11-65
Control
Cineole
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
df
t Stat
Observations
Pooled Variance
5.062215184
169
df
9.55325819
2.784810127
3.27027027
Variance
4.504381694
4.583487597
79
74
Observations
Pooled Variance
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
1.45934E-17
2.77298E-13
-1.407938969
0.161204561
Control
Cineole
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
Pooled Variance
df
Observations
83.12429112
184
43.13829787
51
Variance
79.77636696
84.68817204
94
94
Observations
Pooled Variance
df
Camphor
Mean
82.2322695
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
3.03696E-09
0.795050158
-5.943528856
1.9727986
1.35028E-08
A11-66
Length
Syzygium leuhmannii - Radical
Radicle
Length
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
Mean
Variance
Camphor
48.74468085
Mean
101.908624
125.5470144
Variance
92
94
113.8562887
184
Cineole
52.61956522
Observations
Pooled Variance
df
48.74468085
64.7994738
125.5470144
94
94
Observations
Pooled Variance
df
Camphor
43.26595745
95.17324411
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
4.78229E-11
0.014183313
-3.850089877
1.9727986
0.000162279
Control
Cineole
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
df
1.548087268
184
t Stat
Observations
Pooled Variance
df
4.44798114
1.829158087
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
1.49736E-05
0.075781132
1.9727986
0.028258098
A11-67
Control
Cineole
Control
Mean
87.34782609
80.76595745
Variance
955.2183469
922.912377
92
94
Observations
Pooled Variance
938.889786
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
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
0.144714279
*No sig dif exists in leaf area between Control and Cineole
0.553504934
*No sig dif exists in leaf area between Control and Camphor
1.9727986
0.418889226
*No sig dif exists in leaf area between Cineole and Camphor
Control
Cineole
Control
Mean
43.23913043
37.65116279
Variance
140.2718586
132.276881
92
86
Observations
Pooled Variance
136.4106478
Camphor
43.23913043 31.70212766
Mean
Variance
140.2718586 187.9748341
Variance
Observations
92
94
164.3826017
176
Cineole
Mean
Pooled Variance
df
df
31.70212766
132.276881
187.9748341
86
94
Observations
Pooled Variance
161.377497
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
0.001685787
5.07533E-09
0.001989005
* 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
Control
Cineole
Control
Mean
38.09782609
32
Variance
90.57274247
104.2352941
92
86
Observations
Pooled Variance
97.17113389
t Stat
Mean
Variance
90.57274247 177.9720888
Variance
Observations
92
Observations
Pooled Variance
df
4.12419828
94
134.7474121
176
Cineole
38.09782609 29.53191489
Pooled Variance
df
Camphor
Mean
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
5.72715E-05
1.15103E-06
A11-68
0.167997219
* 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
Control
Cineole
Control
Mean
3.760869565
3.11627907
Variance
3.766364071
3.256908345
92
86
Observations
Pooled Variance
3.520320112
Cineole
3.760869565 1.957446809
Mean
Variance
3.766364071 4.213223519
Variance
Observations
92
94
3.992222379
176
Camphor
Mean
Pooled Variance
df
df
1.957446809
3.256908345
4.213223519
86
94
Observations
Pooled Variance
3.75655616
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
0.023179747
4.60159E-09
4.006844922
9.03311E-05
*Sig.difference exists in leaf No. between the Control and Camphor treatments
*Sig.difference exists in leaf No. between the Cineole and Camphor treatments
A11-69
*Sig.difference exists in leaf No. between the Control and Cineole treatments
*Mean leaf No. is reduced in Cineole treatment
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
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
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
0.089512914
3.70834E-05
0.009472686
* Mean leaf area is sig. reduced by Camphor between Cineole And Camphor treatments
Control
Cineole
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
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
0.071923133
23.38095238
25.109375
Variance
34.09409065
50.44816468
84
64
Observations
41.15098561
138
df
-3.183898426
t Stat
0.00089808
Camphor
Mean
Pooled Variance
df
P(T<=t) one-tail
64
37.77439097
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
0.001796161
0.106554774
A11-70
Control
Mean
Variance
Control
14.82142857
Mean
80.18666667
59.81110155
Variance
76
84
Observations
Pooled Variance
Cineole
19
69.48304702
df
76
64
73.5234375
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
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
0.001852065
8.52488E-05
1.307236171
0.09659376
0.193187521
* 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.
Control
Cineole
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
t Stat
Pooled Variance
df
Observations
0.687285469
148
-3.146838292
1.540540541
0.90625
Variance
0.553128471
0.75297619
74
64
Observations
Pooled Variance
df
0.645704988
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
0.001995986
0.092212799
4.624209138
8.65238E-06
* Sg.dif in leaf no. exists between the Cineole and Camphor treatments.
A11-71
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
138
Camphor
Mean
Pooled Variance
df
-1.008246735
64
176.7356801
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
0.314877605
0.049831855
0.000290851
Control
Mean
Control
40.25510204
33.37096774
Mean
136.748685
183.2863564
Variance
98
62
Variance
Observations
Pooled Variance
Cineole
154.7157607
Cineole
Camphor
44.55932203
Mean
33.37096774
44.55932203
136.748685
224.7679719
Variance
183.2863564
224.7679719
98
59
62
59
169.6849343
158
Camphor
40.25510204
Observations
Pooled Variance
df
Pooled Variance
df
Observations
203.5042867
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
0.000822979
0.046680115
1.65775873
3.35208E-05
A11-72
Control
Mean
Variance
Observations
Pooled Variance
Cineole
11.08064516
Mean
152.128866
80.69830777
Variance
98
62
124.5512454
Control
18.07142857
Observations
Pooled Variance
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
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
0.00016485
0.000128946
-0.025391793
0.489892494
1.65775873
0.979784988
*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.
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' treatment below or above the mean of the 'Control',
and the second symbol identifies significant effect direction in the 'Camphor' treatment.
= : equal significance between treatments.
A12-4
Table A12.2 Summary effect table for t-tests of post-emergent growth in the herbs.
Species
Treatment Significance to
Control.
Growth Significance to
Control
Significance
Between
Treatments.
Mean
Comparison
Cineole to
Camphor
<
<
<
<
Alpinia caerulea
Radicle Length
Ci
Leaf No.
Ci
Leaf Area
Ci Ca
-+
na2
na2
na2
>
>
=
>
Shoot Length
*Chloris gayana
Ci
Cyperus enervis
Shoot Length
Ca
Radicle Length
Ca
Leaf No.
Ca
Leaf Area
Ca
ns
ns
Radicle Length
Ca
Leaf No.
ns
ns
Leaf Area
Ca
=
=
=
=
----
>
=
>
>
--+
=
<
Helichrysum bracteatum
Shoot Length
Lomandra longifolia
Shoot Length
Radicle Length
Ci Ca
Ca
Leaf No.
Ci Ca
Leaf Area
Ci Ca
Oplismenus aemulus
Shoot Length
Ci Ca
Radicle Length
Ci Ca
A12-5
Ci Ca
--
<
=
na2
na2
na2
Ci Ca
--
ns
ns
Leaf No.
Ci
Leaf Area
Ottochloa gracillima
*Paspalum dilatatum
Leaf No.
Ci Ca
--
Leaf Area
ns
ns
=
=
>
=
-+
--
<
<
=
<
Ci Ca
--
Radicle Length
ns
ns
Leaf No.
ns
ns
Leaf Area
ns
ns
>
=
=
=
na2
na2
na2
-+
---
<
=
=
=
Shoot Length
Radicle Length
*Pennisetum clandestinum
Shoot Length
Radicle Length
Ci Ca
Ca
Leaf No.
Ci Ca
Leaf Area
Ci
Pratia purpurescens
Shoot Length
Ci Ca
Radicle Length
Ci Ca
Leaf No.
Ca
Leaf Area
Ci Ca
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' treatment below or above the mean of the 'Control',
and the second symbol identifies significant effect direction in the 'Camphor' treatment.
= : equal significance between treatments.
< : 'Cineole' treatment significantly reduced below 'Camphor' treatment.
> : 'Cineole' treatment significantly increased above 'Camphor' treatment.
2
A12-6
Table A12.3 Summary effect table for t-tests of post-emergent growth in the vines.
Species
Treatment Significance
to Control.
Growth Significance to
Control
Mean Increased or
Decreased
Significance
Between
Treatments.
Mean
Comparison
Cineole to Camphor
* Anredera cordifolia
Shoot Length
Ci
Radicle Length
Ca
Leaf No.
ns
ns
Leaf Area
ns
ns
=
=
=
=
* Cardiospermum grandiflorum
Ci Ca
+
++
Leaf No.
ns
ns
Leaf Area
Ca
<
=
=
=
=
>
=
=
na1
Shoot Length
Radicle Length
Ca
Cissus hypoglauca
Leaf No.
Ci Ca
Leaf Area
Ci Ca
----
na1
na1
Shoot Length
Radicle Length
Geitonoplesium cymosum
Ci Ca
Ca
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' treatment below or above the mean of the 'Control',
and the second symbol identifies significant effect direction in the 'Camphor' treatment.
= : equal significance between treatments.
< : 'Cineole' treatment significantly reduced below 'Camphor' treatment.
> : 'Cineole' treatment significantly increased above 'Camphor' treatment.
1
80
% germination
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 64 67 70 73 76
Days
Figure
13.1a
Figure
1a. Comparison of actual and fitted germination for the
'control' treatment of algae culture 1.
100
90
2
r = 0.972965
80
% germination
70
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 53 55 57 59 61 63 65 67 69 71
Days
Figure
13.1b
Figure
1b. Comparison of actual and fitted germination for the
'cineole' treatment of algae culture 1.
100
observed germination
fitted germination
90
80
% germination
70
60
50
40
30
2
r = 0.984483
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 76
Days
Figure
13.1c
Figure
1c. Comparison of actual and fitted germination for the
'camphor' treatment of algae culture 1.
100
90
r2 = 0.982132
80
% germination
70
60
50
40
30
20
observed germination
fitted germination
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
Days
Figure
2a. Comparison of actual and fitted germination for the 'control'
Figure
13.2a
treatment of algae culture 2.
100
90
80
r2 = 0.985068
% germination
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 64 67 70 73 76
Days
Figure
2b. Comparison of actual and fitted germination for the
Figure
13.2b
100
90
80
% germination
70
60
r2 = 0.997884
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 64 67 70 73
Days
Figure
2c. Comparison of actual and fitted germination for the
Figure
13.2c
A13-2
100
r2 = 0.994844
90
80
% germination
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 64 67 70 73
Days
Figure
Figure
13.3a3a. Comparison of actual and fitted germination for the
100
90
80
% germination
70
r2 = 0.988586
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 64 67 70 73
Days
Figure
Figure
13.3b3b. Comparison of actual and fitted germination for the
100
90
80
% germination
70
60
50
r2 = 0.994882
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 64 67 70 73
Days
Figure
13.3c 3c. Comparison of actual and fitted germination for the
Figure
A13-3
Canopy
Type
Foliage Projective
Cover (%)
Slope
()
Aspect
Elevation
(m)
Soil
Order
Soil
Colour
Soil
pH
Litter
Depth
(mm)
Root
Competition
Animal
Browsing
Other
Disturbance
1.1
camphor
70
12
North
52.9
Ferrasol
2.5YR 2.5/1
5.5
50
100-60
Highly compacted
Absent
Mowing
1.2
camphor
70
12
North
52.9
Ferrasol
2.5YR 2.5/1
5.5
50
100-60
Highly compacted
Absent
Mowing
1.3
camphor
70
12
North
52.9
Ferrasol
2.5YR 2.5/1
5.5
50
100-60
Highly compacted
Absent
Mowing
2.1
control
70
12
North
31.9
Ferrasol
2.5YR 2.5/4
5.5
50
100-60
Highly compacted
Absent
Mowing
2.2
control
70
12
North
31.9
Ferrasol
2.5YR 2.5/4
5.5
50
100-60
Highly compacted
Absent
Mowing
2.3
control
70
12
North
31.9
Ferrasol
2.5YR 2.5/4
5.5
50
100-60
Highly compacted
Absent
Mowing
3.1
control
80
North
47.2
Ferrasol
2.5YR 2.5/3
5.5
50
100-60
Highly compacted
Absent
Mowing
3.2
control
80
North
47.2
Ferrasol
2.5YR 2.5/3
5.5
50
60-40
Highly compacted
Absent
Mowing
3.3
control
80
North
47.2
Ferrasol
2.5YR 2.5/3
5.5
50
100-60
Highly compacted
Absent
Mowing
4.1
camphor
80
North
28.5
Ferrasol
2.5YR 2.5/1
5.5
30
100-60
Highly compacted
Absent
Mowing
4.2
camphor
80
North
28.5
Ferrasol
2.5YR 2.5/1
5.5
30
100-60
Highly compacted
Absent
Mowing
4.3
camphor
80
North
28.5
Ferrasol
2.5YR 2.5/1
5.5
30
100-60
Highly compacted
Absent
Mowing
5.1
control
60
North
30
Ferrasol
2.5YR 2.5/1
5.5
20
60-40
Highly compacted
Absent
Mowing
5.2
control
60
North
30
Ferrasol
2.5YR 2.5/1
5.5
30
100-60
Highly compacted
Absent
Mowing
5.3
control
60
North
30
Ferrasol
2.5YR 2.5/1
5.5
30
100-60
Highly compacted
Absent
Mowing
Site /
Quadrat
No.
Litter
Spread
(%)
A14-2
Table A14.1 Environmental attributes across the sites and quadrats for *C camphora seedlings recorded along a ridge-line at Rosebank, NSW.
Table A14.2 Environmental attributes across the sites and quadrats for *C camphora seedlings recorded along a creek-line at Nimbin, NSW.
Canopy
Type
Foliage Projective
Cover (%)
Slope
()
Aspect
Elevation
(m)
Soil
Order
Soil
Colour
Soil
pH
Litter
Depth
(mm)
Litter
Spread
(%)
Root
Competition
Animal
Browsing
Other
Disturbance
1.1
camphor
80
South
4.3
Podosol
10YR 2/1
7.2
60
100-60
Highly compacted
Absent
Nil
1.2
camphor
80
South
4.3
Podosol
10YR 2/1
7.2
60
100-60
Highly compacted
Absent
Nil
1.3
camphor
80
South
4.3
Podosol
10YR 2/1
7.2
60
100-60
Highly compacted
Absent
Nil
2.1
cineole
80
South
4.3
Podosol
10YR 2/1
7.2
60
100-60
Highly compacted
Absent
Nil
2.2
cineole
80
South
4.3
Podosol
10YR 2/1
7.2
60
100-60
Highly compacted
Absent
Nil
2.3
cineole
80
South
4.3
Podosol
10YR 2/1
7.2
60
100-60
Highly compacted
Absent
Nil
3.1
control
70
10
Podosol
10YR 2/2
30
60-40
Highly compacted
Absent
Nil
3.2
control
70
10
Podosol
10YR 2/2
30
60-40
Highly compacted
Absent
Nil
3.3
control
70
10
Podosol
10YR 2/2
30
60-40
Highly compacted
Absent
Nil
Table A14.3 Environmental attributes across the sites and quadrats for *C camphora seedlings recorded along a creek-line at Uki, NSW.
Site /
Quadrat
No.
Canopy
Type
Foliage Projective
Cover (%)
Slope
()
Aspect
Elevation
(m)
Soil
Order
Soil
Colour
Soil
pH
Litter
Depth
(mm)
Litter
Spread
(%)
Root
Competition
Animal
Browsing
Other
Disturbance
1.1
camphor
80
East
10
Podosol
10YR 2/1
40
100-60
Highly compacted
Absent
River wash
1.2
camphor
80
East
10
Podosol
10YR 2/1
40
100-60
Highly compacted
Absent
River wash
1.3
camphor
80
East
10
Podosol
10YR 2/1
40
100-60
Highly compacted
Absent
River wash
2.1
camphor
90
East
12
Podosol
10YR 2/1
7.2
50
60-40
Highly compacted
Absent
River wash
2.2
camphor
90
East
12
Podosol
10YR 2/1
7.2
50
60-40
Highly compacted
Absent
River wash
2.3
camphor
80
East
12
Podosol
10YR 2/1
7.2
60
100-60
Highly compacted
Absent
River wash
3.1
cineole
80
East
10
Podosol
10YR 2/1
40
100-60
Highly compacted
Absent
River wash
3.2
cineole
80
East
10
Podosol
10YR 2/1
40
100-60
Highly compacted
Absent
River wash
3.3
cineole
80
East
10
Podosol
10YR 2/1
40
100-60
Highly compacted
Absent
River wash
A14-3
Site /
Quadrat
No.
Table A14.4 Environmental attributes across the sites and quadrats for *L. lucidum seedlings recorded along a ridge-line at Rosebank, NSW.
Foliage Projective
Cover (%)
Slope
()
Aspect
Elevation
(m)
Soil
Order
1.1
camphor
70
12
North
52.9
Ferrasol
1.2
camphor
70
12
North
52.9
1.3
camphor
70
12
North
2.1
control
70
12
2.2
control
70
2.3
control
3.1
Soil
Colour
Soil
pH
Litter
Depth
(mm)
Litter
Spread
(%)
Root
Competition
Animal
Browsing
Other
Disturbance
2.5YR 2.5/1
5.5
50
100-60
Highly compacted
Absent
Mowing
Ferrasol
2.5YR 2.5/1
5.5
50
100-60
Highly compacted
Absent
Mowing
52.9
Ferrasol
2.5YR 2.5/1
5.5
50
100-60
Highly compacted
Absent
Mowing
North
31.9
Ferrasol
2.5YR 2.5/4
5.5
50
100-60
Highly compacted
Absent
Mowing
12
North
31.9
Ferrasol
2.5YR 2.5/4
5.5
50
100-60
Highly compacted
Absent
Mowing
70
12
North
31.9
Ferrasol
2.5YR 2.5/4
5.5
50
100-60
Highly compacted
Absent
Mowing
control
80
North
47.2
Ferrasol
2.5YR 2.5/3
5.5
50
100-60
Highly compacted
Absent
Mowing
3.2
control
80
North
47.2
Ferrasol
2.5YR 2.5/4
5.5
50
60-40
Highly compacted
Absent
Mowing
3.3
control
80
North
47.2
Ferrasol
2.5YR 2.5/3
5.5
50
100-60
Highly compacted
Absent
Mowing
4.1
camphor
80
North
28.5
Ferrasol
2.5YR 2.5/1
5.5
30
100-60
Highly compacted
Absent
Mowing
4.2
camphor
80
North
28.5
Ferrasol
2.5YR 2.5/1
5.5
30
100-60
Highly compacted
Absent
Mowing
4.3
camphor
80
North
28.5
Ferrasol
2.5YR 2.5/1
5.5
30
100-60
Highly compacted
Absent
Mowing
5.1
control
60
North
30
Ferrasol
2.5YR 2.5/1
5.5
20
60-40
Not compacted
Absent
Mowing
6.1
control
60
North
25
Ferrasol
2.5YR 2.5/1
5.5
20
60-40
Not compacted
Absent
Mowing
6.2
control
60
North
25
Ferrasol
2.5YR 2.5/1
5.5
20
60-40
Highly compacted
Absent
Mowing
6.3
control
60
North
25
Ferrasol
2.5YR 2.5/1
5.5
20
60-40
Highly compacted
Absent
Mowing
A14-4
Canopy
Type
Site /
Quadrat
No.
Canopy
Type
Foliage Projective
Cover (%)
Slope
()
Aspect
Elevation
(m)
Soil
Order
Soil
Colour
Soil
pH
Litter
Depth
(mm)
1.1
control
70
South
57.5
Ferrasol
7.5YR 3/2
5.5
20
1.2
control
70
South
57.5
Ferrasol
7.5YR 3/2
5.5
1.3
control
70
South
57.5
Ferrasol
7.5YR 3/2
2.1
camphor
70
South
57.5
Ferrasol
2.2
camphor
70
South
57.5
2.3
camphor
70
South
57.5
Site /
Quadrat
No.
Litter
Spread
(%)
Root
Competition
Animal
Browsing
Other
Disturbance
60-40
Highly compacted
Absent
Mowing
20
60-40
Highly compacted
Absent
Mowing
5.5
20
60-40
Highly compacted
Absent
Mowing
7.5YR 3/2
5.5
30
60-40
Highly compacted
Absent
Mowing
Ferrasol
7.5YR 3/2
5.5
30
60-40
Highly compacted
Absent
Mowing
Ferrasol
7.5YR 3/2
5.5
30
60-40
Highly compacted
Absent
Mowing
A14-5
Table A14.5 Environmental attributes across the sites and quadrats for *L. lucidum seedlings recorded along a ridge-line at Lismore, NSW.
Table A14.6 Environmental attributes across the sites and quadrats for G. semiglauca seedlings recorded along a ridge-line at Rosebank, NSW.
Foliage Projective
Cover (%)
Slope
()
Aspect
Elevation
(m)
Soil
Order
1.1
camphor
70
North
60
Ferrasol
1.2
camphor
70
North
60
1.3
camphor
70
North
2.1
control
70
2.2
control
70
2.3
control
3.1
Soil
Colour
Soil
pH
Litter
Depth
(mm)
2.5YR 2.5/2
5.5
30
Ferrasol
2.5YR 2.5/2
5.5
60
Ferrasol
2.5YR 2.5/1
55
Ferrasol
55
70
control
70
3.2
control
70
3.3
control
70
NorthEast
NorthEast
NorthEast
Litter
Spread
(%)
Root
Competition
Animal
Browsing
Other
Disturbance
100-60
Highly compacted
Nil
Mowing
30
100-60
Highly compacted
Nil
Mowing
5.5
50
100-60
Highly compacted
Nil
Mowing
2.5YR 2.5/4
5.5
40
100-60
Highly compacted
Nil
Mowing
Ferrasol
2.5YR 2.5/4
5.5
40
100-60
Highly compacted
Nil
Mowing
55
Ferrasol
2.5YR 2.5/4
5.5
50
60-40
Highly compacted
Nil
Mowing
48
Ferrasol
2.5YR 2.5/2
5.5
40
60-40
Highly compacted
Nil
Mowing
48
Ferrasol
2.5YR 2.5/4
5.5
50
60-40
Highly compacted
Nil
Mowing
48
Ferrasol
2.5YR 2.5/3
5.5
50
60-40
Highly compacted
Nil
Nil
A14-6
Canopy
Type
Site /
Quadrat
No.
Table A14.7 Environmental attributes across the sites and quadrats for G. semiglauca seedlings recorded along a creek-line at Uki, NSW.
Site /
Quadrat
No.
Canopy
Type
Foliage Projective
Cover (%)
Slope
()
Aspect
Elevation
(m)
Soil
Order
Soil
Colour
Soil
pH
Litter
Depth
(mm)
Litter
Spread
(%)
Root
Competition
Animal
Browsing
Other
Disturbance
1.1
control
80
East
10
Podosol
10YR 2/1
7.2
40
100-60
Highly compacted
Nil
Nil
1.2
control
80
East
10
Podosol
10YR 2/1
7.2
40
100-60
Highly compacted
Nil
Nil
1.3
control
80
East
10
Podosol
10YR 2/1
7.2
40
100-60
Highly compacted
Nil
Nil
2.1
camphor
80
East
12
Podosol
10YR 2/1
50
100-60
Highly compacted
Nil
Nil
2.2
camphor
80
East
12
Podosol
10YR 2/1
50
100-60
Highly compacted
Nil
Nil
2.3
camphor
80
East
12
Podosol
10YR 2/1
50
100-60
Highly compacted
Nil
Nil
Table A14.8 Environmental attributes across the sites and quadrats for O. aemulus recorded along a creek-line at Uki, NSW.
Site /
Quadrat
No.
1.1
Soil
Colour
Soil
pH
Litter
Depth
(mm)
Litter
Spread
(%)
Root
Competition
Animal
Browsing
Other
Disturbance
Podosol
10YR 2/1
6.5
40
100-60
Highly compacted
Nil
Nil
10
Podosol
10YR 2/1
6.5
40
100-60
Highly compacted
Nil
Nil
10
Podosol
10YR 2/1
40
100-60
Highly compacted
Nil
Nil
Canopy
Type
Foliage Projective
Cover (%)
Slope
()
Aspect
Elevation
(m)
control
70
South-
Soil
Order
West
1.2
control
70
SouthWest
1.3
control
70
South-
2.1
camphor
70
10
East
10
Podosol
10YR 2/1
7.2
30
100-60
Highly compacted
Nil
Nil
2.2
camphor
70
10
East
10
Podosol
10YR 2/1
7.2
30
100-60
Highly compacted
Nil
Nil
2.3
camphor
70
10
East
10
Podosol
10YR 2/1
7.2
30
100-60
Highly compacted
Nil
Nil
A14-7
West
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
106.5561798
567.9320462
356
520.4291699
0
1358
1.455514102
0.072879056
1.645976467
0.145758113
1.961712355
P(T<=t) two-tail
0.145758113
Camphor
104.5079681
503.616088
1004
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
106.5561798
567.9320462
356
519.0741318
0
723
-9.170936036
2.40057E-19
1.646963911
4.80115E-19
1.9632505
P(T<=t) two-tail
Cineole
122.0785908
471.9421763
369
4.80115E-19
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Camphor
104.5079681
503.616088
1004
495.1142649
0
1371
-12.97117156
1.11929E-36
1.645965812
2.23858E-36
1.961695761
P(T<=t) two-tail
Cineole
122.0785908
471.9421763
369
2.23858E-36
*No Sig. dif in shoot length exists between the Control and Camphor
* Mean shoot length slightly decreased in Camphor compared to Control
*Sig. dif in shoot length exists between the Control and Cineole
* Mean shoot length Sig. Increased in Cineole compared to Control
7
12.05070423
356
4.729930792
0
1358
20.48391645
8.91263E-82
1.645976467
1.78253E-81
1.961712355
P(T<=t) two-tail
1.78253E-81
Camphor
4.251992032
2.138829527
1004
Control
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
P(T<=t) two-tail
7
12.05070423
356
7.303946594
0
723
7.788249059
1.1801E-14
1.646963911
2.3602E-14
1.9632505
2.3602E-14
Cineole
5.436314363
2.724873336
369
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
P(T<=t) two-tail
Camphor
4.251992032
2.138829527
1004
2.296133774
0
1371
-12.83855352
5.19544E-36
1.645965812
1.03909E-35
1.961695761
1.03909E-35
*Sig. dif in leaf No. exists between the Control and Camphor
* Mean leaf No. Sig. decreased in Camphor compared to Control
*Sig. dif in leaf No. exists between the Control and Cineole
* Mean leaf No. Sig. decreased in Cineole compared to Control
Cineole
5.436314363
2.724873336
369
A15-2
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
823.5505618
344473.1495
356
180769.6099
0
1358
5.640138374
1.03235E-08
1.645976467
2.0647E-08
1.961712355
P(T<=t) two-tail
Camphor
675.6294821
122828.6761
1004
2.0647E-08
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
P(T<=t) two-tail
Control
823.5505618
344473.1495
356
235249.1549
0
723
2.815368427
0.002502375
1.646963911
0.00500475
1.9632505
Cineole
722.1056911
129883.6165
369
0.00500475
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Camphor
675.6294821
122828.6761
1004
124722.3436
0
1371
-2.161739386
0.015405426
1.645965812
0.030810852
1.961695761
P(T<=t) two-tail
Cineole
722.1056911
129883.6165
369
0.030810852
*Sig. dif in leaf Area exists between the Control and Cineole
* Mean leaf Area Sig. decreased in Cineole compared to Camphor
*Sig. dif in leaf Area exists between the Camphor and Cineole
* Mean leaf Area Sig. decreased in Camphor compared to Cineole
A15-3
Leaf Number
Leaf Area
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
70.44134078
870.2020922
895
724.8336026
0
1570
3.199951321
0.000700989
1.645824757
0.001401977
1.961476082
P(T<=t) two-tail
0.001401977
Camphor
66.05317578
532.5859256
677
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
9.807821229
37.39255621
895
31.23060329
0
1570
6.854393179
5.12856E-12
1.645824757
1.02571E-11
1.961476082
P(T<=t) two-tail
1.02571E-11
Camphor
7.856720827
23.08151172
677
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
284.4011173
76804.1264
895
58760.13694
0
1570
4.001298243
3.29645E-05
1.645824757
6.59291E-05
1.961476082
P(T<=t) two-tail
6.59291E-05
Camphor
234.9970458
34897.2278
677
Leaf Number
Leaf Area
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
94.640625
576.8934547
128
369.5816047
0
325
10.94787789
2.95981E-24
1.649555622
5.91963E-24
1.96729001
P(T<=t) two-tail
5.91963E-24
Camphor
70.79396985
236.6088523
199
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
6.4609375
2.707123524
128
1.90013348
0
325
6.200890903
8.5193E-10
1.649555622
1.70386E-09
1.96729001
P(T<=t) two-tail
1.70386E-09
Camphor
5.492462312
1.382518654
199
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
255.6328125
32553.65151
128
18262.23655
0
325
8.6484845
1.21694E-16
1.649555622
2.43389E-16
1.96729001
P(T<=t) two-tail
2.43389E-16
Camphor
123.2110553
9095.520887
199
A15-4
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
P(T<=t) two-tail
Control
18.73333333
21.42727273
45
36.68066104
0
79
-6.863092572
6.81916E-10
1.66437141
1.36383E-09
1.990450177
Camphor
28.02777778
55.85634921
36
1.36383E-09
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
8.933333333
2.836363636
45
3.207032349
0
79
5.79918688
6.56479E-08
1.66437141
1.31296E-07
1.990450177
P(T<=t) two-tail
1.31296E-07
Camphor
6.611111111
3.673015873
36
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
P(T<=t) two-tail
Control
103.6666667
1545.227273
45
2384.594585
0
79
-4.566355017
8.99869E-06
1.66437141
1.79974E-05
1.990450177
1.79974E-05
*Sig. Dif in internode lengths exist between the Control and Camphor
*Mean internode length sig. decreased in Control below Camphor
*Sig. Dif in leaf No. exists between the Control and Camphor
*Mean leaf No. sig. decreased in Camphor below Control
*Sig. Dif in leaf area exists between the Control and Camphor
*Mean leaf area sig. decreased in Control below Camphor
Camphor
153.5277778
3439.799206
36
Leaf Number
Leaf Area
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
71.16292135
981.1505639
1068
764.432158
0
1978
4.47328937
4.06989E-06
1.645624349
8.13978E-06
1.961163984
P(T<=t) two-tail
8.13978E-06
Camphor
65.58662281
510.6028066
912
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
9.369850187
32.95305452
1068
26.31108485
0
1978
8.985970105
2.91794E-19
1.645624349
5.83588E-19
1.961163984
P(T<=t) two-tail
5.83588E-19
Camphor
7.291666667
18.53174168
912
Mean
Variance
Observations
Pooled Variance
Hypothesized Mean Difference
df
t Stat
P(T<=t) one-tail
t Critical one-tail
P(T<=t) two-tail
t Critical two-tail
Control
273.338015
69644.17524
1068
51492.77846
0
1978
6.445905368
7.1988E-11
1.645624349
1.43976E-10
1.961163984
P(T<=t) two-tail
1.43976E-10
Camphor
207.3892544
30233.12932
912
A15-5
All Samples Species excluding *C. camphora Shoot & Internode Length