The Ecology of Insect Pests and Fungal Pathogens of Drought

The Ecology of Insect Pests and Fungal Pathogens of

Drought Stressed Eucalypt Plantations in Southern

Queensland

Gilbert Whyte

A Thesis presented to the School of Biology and

Biotechnology Murdoch University Western Australia for

the fulfilment of the requirements of a PhD

April 2012

ii

Declaration

I declare that this thesis is my own account of research and contains as its main content

work that has not previously been submitted for a degree at any tertiary education

institution

Gilbert Whyte

April 2012

iii

Acknowledgments

Thanks to Plantations International Great Southern Plantations and the Australian

Research Council (ARC) for funding this project A special thanks to the foresters at

East Coast Tree Farms for their helpful assistance in the field

Thanks to Dr Paul Barber for his help with the taxonomy of fungal species

Thanks to Dr Monique Sakalidis Dr Vera Andjic Kate Taylor and Francisco Tovar for

their company in the lab

Thanks to my supervisors Dr Treena Burgess and Prof Giles Hardy for their ongoing

encouragement throughout the project I am especially grateful for their patience during

my final years

Thanks to Dr Graham OrsquoHara and Linda Knight for their support

Thanks to Mam Dad brothers and sisters for encouraging me in my pursuits

The most sincere thanks to Suzanne for giving me the greatest love and

encouragement

Dedicated to my two girls Bella and Lizzy

iv

INDEX

1 AN INTRODUCTION TO EUCALYPT PLANTATIONS IN SOUTHERN

QUEENSLAND AND A DISCUSSION OF THE INFLUENCE OF MOISTURE

STRESS ON PEST AND PATHOGEN SUSCEPTIBILITY 1

Plantation Forestry in Australia 1

Traditional Eucalypt Plantation Species 2

New Eucalypt Plantation Species 4

The Queensland Plantation Industry 5

Problems in Plantations 7

Defining Stress 8

Eucalypts and Stress 8

Stress and Insect Pests 9

Stress and Pathogens 17

Plantation Industry Questions 24

Thesis Chapters 24

2 AN INVENTORY OF INSECT SPECIES ASSOCIATED WITH EUCALYPT

PLANTATIONS IN SOUTHERN QUEENSLAND 26

Introduction 26

Materials and Methods 30

Results 32

Longicorn Beetles 34

Eucalypt Weevils 37

Chrysomelid Beetles 40

Giant Wood Moths 45

Case Moths 48

Cup Moths 50

Leaf Bag Worms 53

Eucalypt Leafrollers 56

Leaf Blister Sawfly 59

Eucalypt Gall Wasps 61

Mirid Bugs 63

v

Brown Scale Insects 65

Green Vegetable Bugs 68

Psyllids 70

Leafhoppers 73

Planthoppers 75

Clown Bugs 77

Assassin Bugs 79

Ladybird Beetles 81

Praying Mantids 83

Lacewings 85

Discussion 87

3 AN INVENTORY OF FUNGAL SPECIES ASSOCIATED WITH EUCALYPT


Introduction 91


Results 101

Readeriella epicoccoides 105

Mycosphaerella heimii 108

Mycosphaerella lateralis 111

Mycosphaerella marksii 113

Readeriella eucalypti 117

New Fungal Species 119

Teratosphaeria aurantia sp nov 122

Teratosphaeria biformis sp nov 125

Teratosphaeria micromaculata sp nov 127

Discussion 130

4 PESTS AND PATHOGENS OF EUCALYPT PLANTATIONS IN SOUTHERN

QUEENSLAND EFFECTS OF PLANTATION AGE LOCAL CLIMATE AND

SEASON 134

Introduction 134

Materials and Methodology 138

vi

Results 153

Discussion 184

5 PESTS AND PATHOGENS OF EUCALYPTS AND HYBRIDS A GROWTH

PERFORMANCE TRIAL IN SOUTHERN QUEENSLAND 191

Introduction 191


Results 198

Discussion 216

6 THE PATHOGENICITY OF FUNGI ASSOCIATED WITH STEM BASAL

CANKERS OF EUCALYPT PLANTATIONS 221

Introduction 221


Results 231

Discussion 237

7 GENERAL DISCUSSION 242

Important Pests 242

Important Pathogens 243

Economic Impacts 244

Pest and Pathogen Management 244

General Plantation Health 245

Tree Decline Conceptual Models 246

Limitations of the Study 254

Future Research 257

Concluding Remarks 258

8 REFERENCES 259

vii

Abstract

The eucalypt plantation industry is rapidly expanding to supply an increasing demand

for wood both in Australia and other parts of the world Despite rapid industry

development most eucalypt plantations are restricted to four coastal areas These

include the southwest corner of Western Australia eastern New South Wales the

lsquoGreen Trianglersquo (western Victoria and eastern South Australia) and Tasmania Eucalypt

plantations are traditionally grown in these areas because they have favourable climatic

conditions which allow high productivity Eucalyptus globulus is a fast growing eucalypt

species and is currently the most widely planted species in Australia (553 of all

eucalypt plantations)

More recently plantations have been grown in other parts of Australia which are less

suited to E globulus The eucalypt plantation industry in southern Queensland is in its

infancy and has received less attention from researchers compared with Australiarsquos

main plantation centres Species selection has been a major focus and E dunnii is

quickly emerging as one of the most widely planted species Most of the biological

research of Eucalyptus dunnii has been carried out in plantations in Brazil and South

Africa where the species is an important source of pulp for paper production The

suitability of E dunnii in Australian plantations is still being explored and little is currently

known about its susceptibility to pests pathogens or climatic extremes

This is the first comprehensive study of E dunnii plantations in southern Queensland

Unlike most research in plantations which examines the impacts of insect pests and

fungal pathogens as separate areas of research this study focuses on both groups

simultaneously Understanding the ecology of pests and pathogens is an important

aspect of plantation management and is essential to the development of the plantation

viii

industry in southern Queensland

A large diversity of pests and pathogens were identified from E dunnii plantations

during the study Impacts by insects were generally more severe than pathogens

although most pests and pathogens were found to cause low levels of damage Severe

impacts were caused by chrysomelid beetles such as Paropsisterna cloelia which was

the most destructive chrysomelid species Differences in the abundances of

chrysomelid damage were observed in different aged plantations and between

plantations occurring in different regions of southern Queensland

Several genera of pathogenic fungi were identified and the most abundant species

belonged to the genera Mycosphaerella and Teratosphaeria The most damaging of

these species was Mycosphaerella heimii which was previously unknown in Australia

Canker pathogens such as Holocryphia eucalypti Neofusicoccum ribis and Cytospora

eucalypticola were also common in younger plantations (aged 1-2 years) H eucalypti

was identified as the causal pathogen of lsquosudden death syndromersquo and was the only

pathogen observed to be capable of killing its host

Three new species of foliar fungal pathogens were identified belonging to the genus

Teratosphaeria These species were associated with drought stressed hosts and are

likely to represent a small fraction of a potentially larger assemblage of undescribed

species awaiting discovery in southern Queensland

It was expected that the subtropical climate in southern Queensland would be

conducive to a large diversity of pests and pathogens Despite these expectations

widespread drought in eastern Australia (2003-2007) created atypical conditions within

the region which had adverse effects on many species Drought effects may have

benefited some insects such as those which feed on new foliage produced by stressed

trees however most foliar pathogens appeared to be adversely affected Some canker

ix

pathogens appeared to exploit stressed trees and thirteen weak opportunistic

pathogens were identified from stem cankers and necrotic stem tissues Some

saprophytic fungi may have benefited from greater availability of dead tissue due to a

higher incidence of wilting and premature leaf loss

Although drought effects may have overshadowed the effects of pests and pathogens

the resulting conditions provided valuable insight into the ecology of drought stress in

plantations A conceptual model called the lsquoRecovery-Decline Seesawrsquo has been

developed to illustrate the complex interactions of drought stressed trees and their

associated pests and pathogens The study also contributes valuable information which

aims to facilitate development of the southern Queensland plantation industry

1

1 An Introduction to Eucalypt Plantations in Southern Queensland and a Discussion of the Influence of Moisture Stress on Pest and Pathogen Susceptibility

Plantation Forestry in Australia

Trees belonging to the genus Eucalyptus (LHeacuteritier) are the worldrsquos most planted

hardwood species Eucalypt plantations are widely planted in Europe North America

South America Africa and Southeast Asia The global estimate for eucalypt plantations

is approximately 19 million hectares with India being the largest contributor (8 million

hectares) (Bailey and Duncanson 1998 FAO 2010) Australia has an estimated forested

area of 150 million hectares This vegetation occupies approximately 21 of the

continent and is made up of approximately 1474 million hectares of native forest and

almost 2 million hectares of plantations Approximately half of the plantation area is

comprised of eucalypt species (ABARE 2009 Gavran and Parsons 2011)

There are over 800 species of eucalypts and all but 12 are endemic to Australia

(Brooker and Kleinig 1990 Harden 1991) Extensive clear felling of Australiarsquos native

forests has occurred since European settlement however increasing environmental

awareness has led to the abandonment of unsustainable forestry practises in many

regions (Strozaker et al 2000 Zhang et al 2003) Plantation forestry provides a source

of wood products and helps to relieve pressure on native forests (Anderson 1990

Attiwill 1994 Bell 1999) Afforestation also utilises atmospheric carbon dioxide and

plantations are increasingly being recognised as a carbon sink investment (Winjum et

al 1993 Schimel et al 2001) With the expansion of the plantation industry in Australia

the National Plantation Strategy (1997) has a target of trebling the current plantation

area by 2020 This would mean establishing a further 3 million ha of plantations (80 000

2

ha increase each year) (BRS 1998 Strauss 2001 NAFI 2006 National Forestry

Inventory 2007)

Traditional Eucalypt Plantation Species

Eucalypt plantations are usually harvested in short rotations (7-12 years) so the majority

of species are fast growing Eucalypt species vary greatly in form but only a few species

are selected for their desirable wood properties and rapid growth rate (Nichol et al

1992a) Rapid growth in plantations is heavily dependent on favourable site conditions

(Hillis and Brown 1987 Blake 1990 Eldridge et al 1994 Florence 1996)

Eucalypt plantations are traditionally grown in areas with high rainfall and deep fertile

soils (Blake et al 1990) The two most widely planted eucalypt species in Australia are

E globulus (Blue gum) and E nitens (Shining gum) E globulus is native to Tasmania

the Bass Strait Islands and south-eastern Victoria (Eldridge et al 1994) It is a fast

growing species and produces wood which is suitable for both structural timber and pulp

fibre (Eldridge et al 1994 Bailey and Duncanson 1998) E globulus generally requires

deep fertile soils and high annual rainfall (gt600 mm) for rapid growth (Cotterill et al

1985 Tibbits 1986 National Forestry Inventory 2007) Selective breeding has also

increased the growth rate of E globulus in suboptimal conditions (Eldridge et al 1994

Barbour 1997) For example in south-eastern Victoria E globulus is now grown on a

range of soil types such as gradational clay loams and uniform sands These sites also

vary in annual rainfall from 650 to 1000 mm (Weston et al 1991) Although E globulus

is presently the most widely planted eucalypt species it also has undesirable

characteristics such as sensitivity to frost when exposed to temperatures below -6 degC

For this reason E globulus is generally grown in frost free low altitude sites (Volker et

al 1994 Beadle et al 1996)

3

E nitens (shining gum) has a similar growth rate to E globulus and also prefers high

annual rainfall and deep fertile soils (Eldridge et al 1994) E nitens has a greater

tolerance to frost than E globulus and is the preferred species for plantations in colder

climates such as Tasmania and New Zealand In many overseas countries such as

Chile E globulus is also being replaced by E nitens due to its greater suitability

(Lanfranco and Dungey 2001)

The majority of plantations in Australia are concentrated in four main centres (Figure 1-

1) These include Tasmania coastal New South Wales south-west Western Australia

and the lsquoGreen Trianglersquo (an area spanning the boarder of south-west Victoria and

South Australia) E globulus and E nitens are the two most widely planted species in

these areas More recently plantations have also been established further north on the

east coast of southern Queensland

Figure 1-1 Australiarsquos four main eucalypt plantation centres (red) and the newer Queensland plantation area (blue)

4

New Eucalypt Plantation Species

Australia is essentially an arid continent and most of the interior receives less than

600mm of rain per annum (Sands et al 1999) In the past these areas have been

considered unsuitable for E globulus and E nitens however commercial plantations

are now expanding into these regions using new eucalypt species (Loxton and Forster

2000) E dunnii and E grandis are native to eastern Australia and are now being grown

in southern Queensland where annual rainfall is low to moderate (lt600mm) Due to low

rainfall E globulus and E nitens are often considered to be unsuited to this region

The natural distribution of E dunnii consists of two relatively small populations in

northern New South Wales (Boland et al 1984 Benson and Hager 1993 Specht et al

1995) (Figure 1-2) These populations are estimated to occupy less than 80000 ha and

E dunnii is currently listed as a threatened species under the NSW Threatened Species

Conservation Act 1995 (Briggs and Leigh 1988) E dunnii prefers fertile basaltic and

alluvial soils on the margins of rainforests on north western slopes but it will also grow

on a range of aspects within an altitudinal range of 400m and 650m (Booth et al 1989

Benson and Hagar 1993 Jovanovic et al 2000) The wood fibre of E dunnii is good for

pulp light construction timber and veneers (FAO 1988 2000 Hillis and Brown 1987

Benson and Hager 1993) Genetic variability between different families of E dunnii

suggests there is also potential for improvement through selective breeding (Blake

1990 Arnold et al 1998) Jovanovic et al (2000) used climatic data and interpolation

methods to estimate the potential distribution of E dunnii in Australia China Central

America and South America It was found that E dunnii could potentially grow well in

most of eastern Australia (mainly coastal areas) large areas across southern China

(Manion and Zhang 1989) and numerous localities in Central America and South

5

America Jovanovic et al (2000) also stated that the growth rate of E dunnii would

probably vary between areas depending on climatic conditions

The natural distribution of E grandis is much larger than E dunnii and extends

throughout coastal New South Wales and southern Queensland (Angel et al 1999

Jovanovic et al 2000 Wang et al 1998) (Figure 1-2) E grandis prefers alluvial or

volcanic loams with good drainage and high annual rainfall (725-3500 mm) (common in

valleys riverbanks and flats) (Burns and Honkala 1990) The wood fibre is good for pulp

and construction timber such as joinery plywood panelling boatbuilding flooring and

posts (Burns and Honkala 1990)

The Queensland Plantation Industry

The plantation industry in southern Queensland is in its infancy compared with other

Australian plantation centres and contains less than 5 of the total eucalypt plantation

area of Australia (192 000 ha) (Gavran and Parsons 2011) Unlike high rainfall areas

where eucalypt plantations are traditionally grown the climate of southern Queensland

is subtropical and experiences high temperatures and summer rainfall which leads to

high humidity Average annual rainfall ranges from 400-600 mm per annum with coastal

areas generally receiving greater rainfall than inland areas

As one of the fastest growing eucalypt species Oliveira (1988) reported that E dunnii

outgrew 31 other species of eucalypts during trials at Tres Barras Santa Catharina in

Brazil at an altitude of 775m During the 90s E dunnii and E grandis were also grown

in performance trials in southern Queensland to determine if they were suitable as

plantation species (Lee et al 2000) When compared with other species such as E

globulus (Blue Gum) E tereticornis (Forest Red Gum) E camaldulensis (River Red

6

Gum) and E urophylla (no common name) it was found that E dunnii and E grandis

were generally superior in their rate of growth and wood fibre quality This led to the

establishment of large scale plantations of E dunnii and E grandis in southern

Queensland (Figure 1-2)

Although E grandis and E dunnii were originally planted at similar densities from 1999-

2003 observation after 3-4 years indicated that E dunnii was generally performing

better than E grandis Although E dunnii was originally thought to be more prone to

insect attack (especially psyllids) than E grandis E dunnii was later found to be more

frost resistant which led to greater survival rates during winter (Nixon and Hagedorn

1984 Manion and Zhang 1989 Benson and Hager 1993 Wang et al 1998) E dunnii is

Grafton

Bundaberg

Brisbane

Rockhampton

Coffs Harbour

Urbenville

Natural range of E dunnii

Natural range of E grandis

Eucalypt Plantations

QLD-NSW

Border

Figure 1-2 Eucalypt plantations in southern Queensland (dark blue) and the natural distributions of E dunnii (red) and E grandis (light blue)

7

also less prone to termite attack (Macrotermes natalensis) has a faster growth rate and

has superior wood properties to many eucalypt species (Pereira et al 1986 Ferreira et

al 1993 Marco and Lopez 1995 Oliveira 1998) Since 2003 E dunnii has become one

of the most dominant eucalypt plantation species in the region

Problems in Plantations

A general deterioration in health of many E dunnii plantations in southern Queensland

was observed in 2003 Examinations revealed a number of causes including severe

impacts by insect pests and fungal pathogens This was unexpected given that new

plantations often experience a pest and disease free period in the early stages of

development sometimes called lsquoThe Honeymoon Periodrsquo (Burgess and Wingfield

2002) Further examination of plantations indicated that trees were severely moisture

stressed due to excessively dry weather It was soon realised this period was the

beginning of a drought which would impact much of eastern Australia from 2001-2007

(The South East Queensland Drought Report 2007) It was thought that moisture stress

was increasing the susceptibility of plantation trees to pests and pathogens

An examination of the literature revealed that the influence of moisture stress on the

susceptibility of tree species to pests and pathogens is well documented Given the

extensiveness of the literature it is not surprising that some authors have conflicting

hypotheses These conflicts appear to depend mainly on the species being examined

and the type of association This literature is relevant to understanding the ecology of

pests and pathogens in southern Queensland and the impacts of drought

8

The Influence of Moisture Stress on the Susceptibility of Tree

Species to Pests and Pathogens

Defining Stress

Plant stress is defined as any unfavourable condition or substance which negatively

affects plant metabolism growth or development (Waring and Price 1988 Larsson

1989 Lichtenthaler 1996) Levitt (1980) separated biological stress or strain into lsquoplastic

strainrsquo and lsquoelastic strainrsquo Plastic strain was defined as causing irreversible structural or

chemical damage to the plant while lsquoelastic strainrsquo was defined as that which is

reversible after removal of the cause The tendency for non-genetic factors to affect the

susceptibility of plants to disease is often called lsquopredispositionrsquo Predisposition is

defined by Yarwood (1959) as lsquoan internal degree of susceptibility resulting from

external causesrsquo

Plant stress may be caused by several factors including physical damage such as that

caused by lsquowind throwrsquo (Brewer and Merritt 1978) extreme weather such as hail (Smith

and Kemp 1994) frost damage (Linnard 1969 Paton 1981 Kozlowski et al 1991 Ball

et al 1997) high temperatures (County and County 2003) nutrient deficiency or low

water availability (Jacobs 1955 Xu and Dell 1997) Moisture stress is the focus of the

current study and is defined as stress caused by a lack of available water such as

drought (Bradford and Hsiao 1982 Bachelard 1986 Beadle 2000)

Eucalypts and Stress

Although many eucalypt species are adapted to survive in dry arid regions most

plantation species are native to high rainfall areas (Jacobs 1955 Weston et al 1991

Madeira et al 2002) A number of structural and physiological adaptations can influence

9

the susceptibility of eucalypts to moisture stress Species with a root system composed

mainly of surface lateral roots are often more susceptible to moisture stress than

species with deep tap roots E camaldulensis for example has good drought resistance

due to the presence of deep tap roots that can reach ground water (Jacobs 1955)

Foliar characteristics may also have an influence on drought resistance For example

E globulus is more susceptible to moisture stress than E nitens because stomatal

conductance of E globulus foliage is slower to respond to moisture stress which allows

higher rates of water loss through transpiration (White 1996) Symptoms of moisture

stress in eucalypts may include stunted canopies stunted root systems malformation of

the bark (cracks swellings or discolouration) premature leaf loss (Orshan 1954) and or

deterioration of foliage (yellowing reddening purpling or necrosis) (Stone and Bacon

1994 Landsberg 1990)

Stress and Insect Pests

There are many reviews which examine the influence of moisture stress on tree hosts

and their associated insect pests (Mattson and Haack 1987 Larsson 1989 Koricheva

and Larsson 1998 Huberty and Denno 2004 Lieutier 2004) Most research describes

two opposing hypotheses which predict different effects These are the lsquoPlant Vigour

Hypothesisrsquo and the lsquoThe Plant Stress Hypothesisrsquo

The lsquoPlant Vigour Hypothesisrsquo predicts that plants with greater vigour are more attractive

to insect herbivores (Price 1991) This is mainly because vigorously growing plants may

have increased resources higher food quality and a lack of defensive compounds

(Price 1991) Inbar et al (2001) showed support for the lsquoPlant Vigour Hypothesisrsquo by

examining the interaction of feeding insects on plants subjected to various levels of

moisture stress Durzan (1974) tested the plant vigour hypothesis and showed that

10

when nitrogen fertiliser was applied to trees their health increased causing the

production of arginine in foliage which is an attractant to insects Support for the lsquoPlant

Vigour Hypothesisrsquo is also evident in that insect pests often prefer the active growing

parts of a plant to feed upon (Price 1991)

Alternatively the lsquoPlant Stress Hypothesisrsquo predicts that stressed plants are more

susceptible to attack by insect herbivores than healthy plants because plant stress can

inhibit the production of antiherbivore chemicals (White 1969 1984 Louda and Collinge

1992 Koricheva and Larsson 1998) Defensive chemicals have a range of effects on

feeding insects (Taylor 1997 Strauss and Agrawal 1999 Sanson et al 2001) Toxic

compounds such as cyanide may act as a direct deterrent to insects while other

compounds such as tannins may bind to nutrients such as nitrogen and reduce their

availability within plant tissues Nitrogen availability is one of the most important factors

in plant-insect associations (Carne 1965 White 1974 Fox and Macauley 1977 McClure

1980 Ohmart et al 1985 Ohmart et al 1987 Cromer and Williams 1982 Kavanagh

and Lambert 1990 Stone and Bacon 1995 Landsberg and Cork 1997) White (1974)

proposed that the availability of soluble nitrogen is greater in senescing plant tissue

which has a positive effect on insect herbivores The majority of research also supports

the Plant Stress Hypothesis (Krauss 1969 White 1984 Adams and Atkinson 1991

Waring and Cobb 1992 Landsberg and Gillieson 1995 Marschner 1995 Landsberg

and Cork 1997 Zanger et al 1997 Koricheva and Larsson 1998)

Some insects such as borers have the ability to select stressed hosts (Cooper 2001)

Once a host is selected pheromones may be excreted by the colonising individual to

attract other individuals The phenomenon known as lsquomass attackrsquo is common amongst

bark beetle species (Scolytidae) (Wood 1982) Although there is less evidence of mass

11

attack in other borer families it has been proposed that borers in the Cerambycidae may

have similar behaviour (Lawson et al 2002) Increased feeding pressure caused by

mass attack can overpower host defences and cause further stress of the host (Carter

1973 Agrios 1980 Hatcher 1995) This can result in a feedback loop mechanism

(Carne 1965 Landsberg 1990a Landsberg 1990b Landsberg 1990c Stone and Bacon

1995 Landsberg and Cork 1997)

Defoliating pests may also benefit from host stress because the foliage produced by

stressed trees may differ in physical and chemical properties to foliage produced by

healthy trees (Day 1998) One of the main differences between stressed and non-

stressed eucalypts is the greater ratio of juvenile to adult leaves that tend to occur in

stressed trees (heteroblasty) Some insects prefer to feed on foliage which is at a

specific stage of development (Larsson and Ohmart 1988) Chrysomelid beetles prefer

soft juvenile regrowth to tougher adult leaves and may feed more intensively on

stressed hosts (Tanton and Khan 1978 Edwards 1982 Edwards and Wanjura 1990)

Stressed trees may be continually defoliated which can lead to dieback or even death

(Landsberg 1990 Larsson and Ohmart 1988) While few studies conflict with the Plant

Stress Hypothesis some authors offer simpler explanations such as that the impact of

pests on stressed hosts may only appear greater due to other effects such as reduced

plant growth (Stone and Bacon 1995)

The effects of host stress on insects may vary depending on the type of association

Larsson (1989) suggested that sap-sucking species would benefit more from feeding on

stressed hosts than defoliating species due to a more intimate association It has also

been suggested that the effects of host stress may differ between borers and defoliators

depending on the level of the stress Moderate host stress may benefit borer species

12

(Lieutier 2002) while defoliator species may benefit more from severe stress if it leads

to increased available nitrogen in foliage (White 1969 White 1986 Mattson and Haak

1987 Larsson 1989 Larsson and BjOumlrkman 1993) However severe stress may

eventually disadvantage borers and defoliators if it leads to poor host quality (Rouault et

al 2006)

There has been little research examining the role of moisture stress on insects that feed

on eucalypts This may indicate that stress in eucalypts has a limited influence on

feeding insects For example some studies show that the antiherbivore chemicals

produced by eucalypt foliage have little influence on patterns of herbivory Fox and

Macauley (1977) showed that tannin and phenol concentrations in eucalypt foliage have

little effect on the growth rate of the leaf beetle Paropsis atomaria These effects have

been similarly illustrated by Morrow and Fox (1980) who showed that the composition of

herbivore assemblages are very similar between eucalypt species with varying

concentrations of essential oils

Several detailed studies of insects and stressed trees involve conifer species in the

northern hemisphere (Edmunds and Alstad 1978) This may be due to several

biogeographical factors for example the Scolytidae contains several species known to

exploit stressed conifer hosts in the northern hemisphere (Paine et al 1987)

A summary of some of the more well-known examples of insects which exploit stressed

tree hosts has been tabulated presenting a range of species from different taxonomic

groups (Table 11)

13

Pest Host range Distribution Symptoms Ecology References

Borers

Ips acuminatus (Scolytidae)

Pinus spp Europe The larvae create radiating feeding tracks as they feed beneath the bark Adult beetles emerge through holes in the bark surface

The beetles overwinter in leaf litter and occasionally under bark They breed in freshly cut pine wood or trimmed branches Males attack trees first and produce a pheromone attractant which draws other beetles Eggs are laid in galleries excavated by adults under tree bark

Beetles can affect healthy hosts but prefer to invade moisture stressed or weakened trees Adult beetles also prefer to attack stems with thinner bark

Gueacuterarda et al 2000

Sauvard 2004

Ips sexdentatus (Scolytidae)

Abies spp

Larix spp

Pinus spp

Picea spp

Asia the Pacific (Mainland) and Europe

The larvae create radiating feeding tracks as they feed beneath the bark Adult beetles emerge through holes in the bark surface

Attacks are initiated by male beetles who construct nuptial chambers under the bark The males secrete pheromones to attract females who mate and lay eggs within the gallery

Beetles can affect healthy hosts but prefer to invade moisture stressed or weakened trees Beetles also occasionally attack freshly felled trees or windthrown trees

Croiseacute and Lieutier 1993

Dobbertin et al 2007

Ips typographus (Scolytidae)

Picea spp Europe The larvae create radiating feeding tracks as they feed beneath the bark Adult beetles emerge through holes in the bark surface

Adult beetles lay eggs in excavations beneath the bark and the resulting larvae develop in the inner bark (feeding on living and dead phloem tissues)

Beetles can affect healthy hosts but prefer to invade the stems of trees which are drought stressed Beetles also produce pheromones to attract other beetles to stressed trees (mass attack) Adult female beetles transmit the fungus Ceratocystis polonica during oviposition There is also evidence that high temperatures increase the success rate of ovipostion by allowing extended periods of flying and may increase the rate of larval development

Christiansen amp Ericsson 1986

Christiansen amp Bakke 1988

Christiansen 1992


Table 11 Insects species which exploit stressed tree species

14

Borers continued


Phoracantha spp (Cerambycidae)

Eucalyptus spp

Acacia spp

Australia and South east Asia

Larvae create tracks as they feed beneath the bark The stem may become dark and strongly discoloured Frass may accumulate around the base of the stem beneath emergence holes The shape of emergence holes can vary between Phoracantha species

The adult beetles lay eggs on the bark and the resulting larvae develop in the inner bark (feeding on living and dead phloem tissues) of trees

Beetles are believed to be attracted to stressed hosts and adult female beetles can detect stressed hosts presumably by the composition of essential oils secreted by the leaves

Duffy 1963

Hanks et al 1999

Lanfranco and Dungey 2001

Griffiths et al 2004

Pissodes strobi (Curculionidae)

Pinus banksiana

Pinus strobes

Picea abies

North America Larvae create irregular tunnels beneath the bark as they feed Adult beetles emerge through emergence holes in the bark surface

Adult beetles lay eggs beneath the bark of trees and the resultant larvae feed on the phloem

Beetles prefer drought stressed hosts which have higher phloem quality and usually select young trees on open sites

Alfaro and Omule 1990

Lavallee 1994

Pityogenes chalcographus (Scolytidae)

Larix decidua

Picea spp

Pinus spp

Europe The larvae create radiating feeding tracks as they feed beneath the bark Adult beetles emerge through holes in the bark surface

Transmission of Ophiostoma spp to the stem may further reduce host defences

Beetles prefer to attack drought stressed trees Stress caused by damage by Ips typographus can also facilitate infestations

Schwerdtfeger 1929

Avtzis et al 2000

Kirisits 2004

Tomicus piniperda (Scolytidae)

Pinus sp

Picea sp

Europe

North West Africa

Northern Asia


The adult beetles lay eggs on the bark and the resulting larvae develop in the inner bark

Unlike most bark beetles the beetles do not use pheromones to attract mates or mass attack Instead the beetles are attracted to resin scent emitted by stressed hosts

Beetles commonly infest windblown trees lying on the ground and fire-killed standing trees

Davies and King 1977

Vasconcelos et al 2003

15

Defoliating Insects


Chrysomelidae Eucalyptus spp Mainly Australia Young larvae often feed in aggregations and devour entire leaves More mature larvae tend to feed on leaf margins towards the midrib in a semicircle (scalping)

Adult beetles may overwinter in leaf litter or beneath bark Eggs are generally laid on tree hosts in spring and the resultant larvae feed on the foliage

Some chrysomelid species prefer soft juvenile foliage rather than tougher mature adult foliage Stressed trees often produce large quantities of soft epicormic regrowth which is exploited by the beetles

Tanton and Khan 1978

Miles et al 1982

Lymantria dispar (Lymantriidae)

Quercus spp

Tsuga canadensis

Europe

Asia

North America

Larvae are defoliators The first instar larvae chew small holes in leaves The second and third instars feed from the outer edge of the leaf toward the centre

Adult moths lay egg masses on branches and trunks of trees (also human dwellings) Newly hatched larvae disperse on silken threads (up to one mile) Larvae feed on hosts they come into contact with

Although neither adult not larvae target drought stressed hosts these trees are more heavily defoliated which may be due to their more palatable foliage or their lower rate of recovery

Miller and Wallner 1989

Davidson et al 1999

Thaumetopoea pityocampa (Thaumetopoeidae)

Cedrus spp

Larix spp

Pinus spp

Europe Larvae live within large communal nests which are spun from silk Large nests may also contain quantities of frass and faecal pellets

Adult moths are attracted to stressed hosts on which they lay their eggs The resultant larvae feed on the stressed trees which have higher available nitrogen in their leaves compared with healthy trees

Hodar and Zamora 2002

Rouault et al 2006

Buffo et al 2007

Tortrix viridana (Tortricidae)

Quercus spp

Acer spp

Betula spp

Fagus spp

Populus spp

Europe First instar larvae bore into new growth including new buds

More developed larvae shelter within rolled leaves which are spun together with silk

Adult moths lay eggs near leaf buds which the larvae consume when they emerge Larvae eat larger leaves as they develop and pupate within rolled leaves

Moths may be attracted to drought stressed hosts Timing of budburst may also influence susceptibility

Gasow 1925

Schwerdtfeger 1971

Larsson et al 2000

Rubtsov and Utkina 2003

16

Other Insects


Psyllidae Eucalyptus spp Australia Nymphs feed by sap-sucking on the surface of foliage A lsquolerprsquo made of sugar is excreted by the insect to conceal it as it feeds on the leaf surface Lerp shape is often characteristic of species

Adult psyllids lay eggs on foliage and stems and the resultant nymphs disperse on foliage

Stressed trees often have higher nitrogen availability in foliage which accelerates the rate of development of nymphs This leads to greater feeding pressure on stressed hosts

White 1969

Miles et al 1982

17

Stress and Pathogens

The interaction between stressed plants and their pathogens has long been recognised

(Yarwood 1959 Hepting 1963 Bertrand et al 1967 Colhoun 1973 Schoeneweiss

1875 1981 Boyer 1995) Several studies show that stressed plants have modified

tissues which can increase susceptibility to pathogens (Plant Stress Hypothesis) This is

mainly because low water availability in plant tissues can modify or inhibit the activity of

enzymes which are important in defence against pathogens (Slatyer 1967 Kramer

1969 Kolattukudy and Koller 1983 Kolattukudy PE 1985 Boyer 1995)

Fungi respond differently to host stress depending on their ecological role and their

pathogenicity Endophytes are fungi which infect healthy hosts in the absence of a

disease response (Carroll and Carroll 1978 Fisher and Petrini 1990 1992 Kendrick

1992 Fisher et al 1993 Carroll 1997 1988 Arnold et al 2000) Some endophytes are

better described as opportunistic pathogens because they can induce a disease

response if their host becomes stressed These pathogens are also referred to as lsquolatent

pathogensrsquo (Anselmi et al 2007) Saprophytic fungi only infect dead tissue however

some opportunistic saprophytes may cause disease in living tissue if the host is

severely stressed (Bier 1959 1961 Griffin 1977 Rayner and Boddy 1988 Bettucci and

Saravay 1993 Bettucci and Alonso 1997 Bettucci et al 1999)

Moisture stress can reduce the rate of recovery of a host after damage has been

inflicted by a pathogen In the United States in the 1940s Pole Blight a disease of

western white pine (Pinus monticola) was found to be severe during drought conditions

An examination of the trees revealed that they were infected with root pathogens

(unknown species) and that drought conditions caused trees to have almost no root

regenerating capacity (Desprez-Loustau et al 2006) Some authors argue that

18

aggressive pathogens will infect their hosts regardless of stress and that the influence of

stress on disease susceptibility is on disease development rather than the probability of

infection (Walker and Stahmann 1955 Cook and Papendick 1972)

Moisture stress may benefit pathogens by increasing host susceptibility however water

availability is also important for pathogen development Many pathogens depend on

high humidity for producing spores (sporulation) and rainfall is also an important agent

of spore dispersal (splash dispersal) (Howe 1955 Walklate et al 1989 Daniel and Shen

1991 Agrios 2005) Leaf wetness has been shown to increase rates of infection by

foliar pathogens (Beaumont 1947 Krausse and Massie 1975) Flooding plant roots with

water has also been shown to predispose plants to infection by pathogens (Stolzy et al

1965 Duniway and Gordon 1986) Tinsley (1953) showed that increasing the availability

of water to plants in the nursery could increase their susceptibility to viruses

Chrysoporthe cubensis is a canker pathogen of eucalypt plantations grown in high

rainfall areas (van Heerden and Wingfield 2002) By measuring cambial lesions on E

grandis seedlings Swart and Conradie (1992) demonstrated that the pathogenicity of C

cubensis was greater on healthy rather than moisture stressed hosts (Plant Vigour

Hypothesis) Similarly Cytospora species have been shown to have a greater

pathogenicity in healthy hosts of Acer saccharum compared with moisture stressed

hosts (McPartland 1983) Similar patterns also occur with Thyronectria species which

cause cankers of honeylocust (Jacobi and Riffle 1989) The effect of host stress on

fungal pathogens depends ultimately on the species involved and their ecology Some

of the more documented examples have been tabulated (Table 12) These examples

come from all over the world and include a range of host species

19

Pathogen Host range Distribution Symptoms Ecology References

Armillaria spp At least 50 families and over 200 species

(Eucalyptus spp amp Acacia spp in Australia)

(Quercus spp in Europe)

Worldwide Symptoms may differ between Armillaria spp May include dieback of the limbs and branches yellowing of foliage splits exudates and scarring of the stem poor vigour kino exudates from the stem darkening of larger roots Removal of the bark may reveal the presence of mycelial fans

Opportunistic pathogens that infect droughtmoisture stressed trees more successfully than healthy trees Stress may be caused by drought or waterlogged soils

Pathogenicity is variable between Armillaria species

A mellea has been shown to only cause disease in stressed Quercus seedlings

Pearce and Malajczuk 1990

Wargo 1996

Metaliaj 2003

Biscognauxia mediterranea

Fagus silvatica

and Quercus spp

Mediterranean Symptoms include cankers and necrosis of the bark (stems and branches)

An endophyte and opportunistic pathogen that will infect stressed trees more successfully than healthy trees Stress may be caused by droughtmoisture stress

Hendry et al 1998

Franceschini et al 2004

Desprez-Loustau et al 2006

Botryosphaeria dothidea Wide range of trees and shrubs

A major problem in planted forests including Eucalyptus spp

Worldwide Symptoms differ between host species May include fruit rots leaf spots and stem cankers

May cause dieback in large forest trees

A latent symptomless endophyte in healthy leaves of Eucalyptus spp (mainly a pathogen of stressed hosts)

Also a saprophyte of several tree species including Birch (Betula alba) May infect the bark of dead stems when the host is in vigorous condition but will infect living tissues and cause cankers in moisture stressed hosts

Barr 1972

Crist and Shoeneweiss 1975

Zhonghua et al 2001

Table 12 Pathogens which exploit stressed tree species

20


Thielaviopsis paradoxa

Phoenix dactylifera

Saudi Arabia

Iraq

Symptoms include trunk rot bud rot and senescing inflorescences

An opportunistic pathogen mainly affecting stressed palms that have been predisposed to droughtmoisture stress

Sporulates on senescing structures

Suleman et al 2001

Paulin-Mahady et al 2002

Cryphonectria parasitica Castanea spp

Quercus spp

Castanopsis spp

Acer spp

Rhus spp

Typhina spp

Carya ovata

Europe

Asia

Africa

North America

Symptoms include stem cankers caused by infection of the vascular cambium This causes disruption of xylem and phloem (girdling) Severely affected hosts may exhibit premature leaf loss and leaf senescence which may lead to death

A latent pathogen during winter which is often expressed in spring as bark lesions

Mainly spread by wind and rain but may also have insect vectors

Lesions develop quicker on moisture stressed hosts Lesions also develop quicker during the warmer months compared with the autumn and winter (rainfall and temperature dependent)

Shear et al 1917

Hepting 1974

Anagnostakis1984

Waldboth and Oberhuber 2009

Cytophoma pruinosa Fraxinus spp North America Symptoms include stem cankers cracking swelling and discolouration of the bark

A latent pathogen of healthy trees which only causes disease symptoms when the host is drought moisture stressed

Ross 1964

Tobiessen and Buchsbaum 1976

Rayner and Boddy 1988

Cryptostroma corticale Acer spp North America

Britain

Symptoms include premature leaf loss and leaf senescence (die-back) premature bark shedding and the production of a thick layer of brownish-black dry phialospores on the bark

An opportunistic pathogen mainly infecting drought stressed trees or trees growing on shallow soils

Gibbs et al 1997

Cytospora chrysosperma Populus tremuloides

Acer spp

Populus spp

North America Symptoms include stem cankers which are generally brownish-yellow sunken areas irregular in outline and range from diffuse to slightly target-shaped The bark may split at canker margins the inner bark turns black and wood beneath the canker is stained brown and water soaked White masses of spores may be produced during wet weather

An opportunistic pathogen mainly infecting drought moisture stressed hosts

May also infect hosts which are stressed due to repeated defoliation by insects

Christensen 1940

Bertrand 1967

Jones 1985

Guyon et al1996

21


Cytospora eucalypticola Eucalyptus spp South Africa

Australia

The main symptom of infection is the presence of small cankers on the bark of stems

A mild pathogen mainly infecting drought moisture stressed hosts

Infected lesions usually heal rapidly although the fungus can persist in the tissues

Shearer et al 1987

Old et al 1990

Diplodia mutila Pinus spp

Quercus spp

Juniperus spp

Fraxinus spp

Eucalyptus spp

Europe

South America

North America

Symptoms include cankers and blackening of the stem

An opportunistic pathogen with a broad host range The species often affects drought moisture stressed hosts

Luque and Girbal 1989

Luque et al 2002

Diplodia pinea A range of Conifer species

Pinus spp

Picea spp

Abies spp

Worldwide Symptoms include shoot blight crown wilt root rot and stem cankers

A saprophyte of dead wood and a mild pathogen affecting drought moisture stressed hosts May also be endophytic

Infection may also be facilitated by wounding such as by hail or pruning

Birch 1937

Laughton1937

Eldridge 1961

Lűckhoff 1964

Buchanan 1967

Marks and Minko 1969

Punithalingham and Waterson 1970

Barker 1979

Gibson 1980

Brown et al 1981

Chou 1982

Swart et al 1985

22


Holocryphia eucalypti Eucalyptus spp

North America

Australia

South Africa

Symptoms include basal stem cankers kino exudation and in severe cases branch and shoot dieback

An opportunistic pathogen often associated with drought moisture stressed hosts

Levels of carbohydrate in the stem may influence susceptibility Moisture stressed trees have lower carbohydrate levels which may facilitate infection

Schoenweiss 1975

Davison 1982

Appel and Stipes 1984

Walker et al 1985

Roane et al 1986

Old et al 1990

Gryzenhout 2006

Gryzenhout et al 2006

Hypoxolon mediterraneum

Quercus spp

Castanea spp

Populus spp

Europe Symptoms include stem cankers cracking of the bark and blackening of the vascular cambium

An opportunistic pathogen mainly affecting drought moisture stressed hosts

H mediterraneum has also been found in association with Cryphonectria parasitica C parasitica may facilitate infection by H mediterraneum by causing cankers which create entry points

Bruck and Manion 1980

Agosteo and Pennisi 1990

Valentini 1994

Neofusicoccum ribis

Various Eucalyptus hosts including

E dunnii

E grandis

E camaldulensis

E radiata

E cladocalyx

E marginata

Corymbia calophylla

Worldwide Symptoms include leaf spotsblights stem cankers sinking and swelling of the stem cracking of the bark and brown streaking of the heartwood

A broad range pathogen often associated with droughtmoisture stressed hosts

Davidson and Tay 1983

Shearer et al 1987

Old et al 1990

Luque et al 2002

Slippers et al 2004

23


Phomopsis alnea Alnus spp Europe

Southeast Asia

Africa

Symptoms include stem cankers which are sunken irregularly to circular shaped water-soaked and with abundant dark exudates

A saprophyte and weak pathogen mainly affecting droughtmoisture stressed hosts

Surico et al1996

Moricca 2002

Septoria musiva Populus spp

Aspen spp

North America

Crimea and the Caucasus region of Asia

Symptoms include leaf spotting (which can lead to defoliation) and stem cankers which often develop on the primary shoots of 2 and 3-year-old poplars Infections may also lead to stem breakage

An opportunistic pathogen mainly affecting droughtmoisture stressed trees

Lower moisture content of host tissues may increase susceptibility to infection

Bier 1939

Thomson 1941

Teterevnikova and Babayan 1976

Ostry and McNabb 1983 and 1986

Moore and Wilson 1983

Abebe and Hart 1990

Maxwell 1997

Xylella fastidiosa

Parthenocissis quinquefolia

and Citrus spp

United States Symptoms include leaf scorching along the stem of Parthenocissis quinquefolia and variegated chlorosis of the foliage of Citrus spp

An opportunistic pathogen mainly affecting droughtmoisture stressed hosts

Boyer 1995

24

Plantation Industry Questions

The plantation industry in southern Queensland is in its infancy Previous pest and

disease management tools have been adopted from other plantation centres but a

greater knowledge of pests and diseases within the region is required to further industry

development

The following questions are relevant to the development of the plantation industry in

southern Queensland

1 How diverse are pests and pathogens in plantations within the southern

Queensland region

2 Where do pests and pathogens originate (native or exotic)

3 Are pests and pathogens affected by the maturation of plantations (effects of

plantation age)

4 Are there differences in the distribution of pests and pathogens within the region

(effects of local climate)

5 Are pests and pathogens affected by seasonal changes (effects of season)

6 How do eucalypt species and their hybrids vary in their susceptibility to pests and

pathogens

7 Can an ecological understanding of important pests or pathogens lead to the

development of better methods of control

Thesis Chapters

In 2003 a three year study of the pests and pathogens of eucalypt plantations in

southern Queensland was commenced The unifying aim of this study was to learn

more about the ecology of pests and pathogens in southern Queensland plantations

25

and to convey this information to the industry This was achieved by field observation

the collection of biological material and by conducting experiments which address each

of the questions previously presented This information has been organised into a series

of chapters and a general discussion

Chapter 2 An Inventory of Insect Species Associated with Eucalypt Plantations in

Southern Queensland

Chapter 3 An Inventory of Fungal Species Associated with Eucalypt Plantations in

Southern Queensland

Chapter 4 Pests and Pathogens of Eucalypt Plantations in Southern Queensland

Effects of Plantation Age Local Climate and Season

Chapter 5 Pests and Pathogens of Eucalypts and their Hybrids A Growth

Performance Trial in Southern Queensland

Chapter 6 Pathogenicity of Fungi Associated with Basal Cankers of Eucalypt

Plantations

Chapter 7 General Discussion

26

2 An Inventory of Insect Species Associated with Eucalypt Plantations in Southern Queensland

Introduction

The genus Eucalyptus is host to hundreds if not thousands of insect herbivores and

commensals (New 1943 Morrow 1977 Ohmart et al 1983a Stone and Bacon 1995)

One study of mixed forest comprising E delegatensis E pauciflora and E dives found

that the density of feeding insects was estimated at 91000 individuals per hectare

(Ohmart et al 1983b) Majer et al (1997) estimated there could be as many as 15000-

20000 phytophagous insect species associated with eucalypt species It is not

surprising therefore that most of Australiarsquos important eucalypt plantation pests are

native (de Little 1989 Abbott et al 1991 Harrington and Ewel 1997 Straus 2001) As

the area of eucalypt plantations has increased in Australia the number of associated

insect pests has also increased (Cooper 2001 Loch and Floyd 2001 Steinbauer 2001

Stone 2001)

Several factors contribute to creating favourable conditions for pests in plantations

Eucalypt species selection is important because species vary in their susceptibility to

insect pests (Macauley and Fox 1980 Richardson and Meakins 1986) Differences in

susceptibility may also occur at a subgeneric level within eucalypts and Monocalyptus

species are generally more susceptible to pests than Symphyomyrtus species (Adams

and Atkinson 1991 Florence 1996 Noble 1989) Pest susceptibility also occurs at an

individual level and several studies show that individual trees of the same species may

respond differently to the same pests This is mainly because of genetic differences

between trees and interactions with the immediate environment (Clark 1962 Carne

1965 1966 Carne et al 1974 Mazanec 1974 Journet 1980)

27

The heritability of pest susceptibility has allowed selective breeding of highly resistant

eucalypt genotypes (Durzan 1974 Altieri and Letourneau 1984 Schowalter et al

1986 Andow 1991 Denison and Kietzka 1993 Laranjeiro 1994 Soria and Borralho

1997) Although selective breeding has improved productivity in plantations by reducing

losses from insect pests it has also led to less desirable effects such as lsquomonoculture

effectsrsquo Root (1973) proposed two explanations for monoculture effects 1) The lsquoNatural

Enemy Hypothesisrsquo - Natural enemies are more effective at controlling pests in diverse

systems rather than monocultures and 2) The lsquoResource Concentration Hypothesisrsquo -

Specialist insect herbivores find it easier to feed and reproduce in monocultures than in

diverse systems Most studies support the Resource Concentration Hypothesis (Jones

and Gibson 1966 Campbell 1972 Rausher 1981 Lawton 1983 Altieri and Letourneau

1984 Schowalter et al 1986 Andow 1991) Monoculture effects may be reduced by

increasing the genetic variability of plantations either by growing different genotypes of

the same species or by growing mixed plant species (Andow 1991 Campbell 1972

Risch 1983 Khanna 1997 Bauhus et al 2000)

Site selection can also influence the risk of infestation by pests For example adult

beetles of some Anoplognathus spp feed on eucalypt foliage while the larvae feed on

the roots of grasses Improving the nutrient content of pasture occurring near

plantations has been shown to benefit the larval stage of the beetle which can lead to

greater numbers of emerging adults that cause damage in nearby plantations (Carne et

al 1974 Urquhart and Stone 1995 Landsberg and Cork 1997) Local climatic

conditions such as high temperature and rainfall may also benefit pests (Howe 1955

Stork 1988 Hill 1994 Nair 2001) Plantations grown in tropical areas may be exposed

to a greater diversity of insect pests than those grown in temperate areas because

insect diversity is typically higher in the tropics (Stork 1988 Speight and Wylie 2001)

28

In established plantations pests may be accidently introduced in association with

germplasm such as seeds seedlings contaminated soil or even land preparation

equipment (Floyd et al 1998) Colonisation of pests from native forests or other

eucalypt plantations may also occur and the proximity of neighbouring trees and the

relatedness of the species can influence the rate of colonisation (Lodge 1993

Harrington and Ewel 1997 Burgess and Wingfield 2002) The lsquoEnemy Release

Hypothesisrsquo describes how plantations which are grown outside their natural range may

be removed from their natural pests The absence of pests may result in greater health

and hence more vigorous growth of plantation trees (Keane and Crawley 2002 Mitchell

and Power 2003 Wingfield 2001)

Due to the growing economic importance of eucalypt plantations in Australia most

entomological research has been aimed at reducing impacts of insect pests (CALM

1990 Turnbull 2000) Some of the more important pests of plantations in Australia

include African black beetle (Heteronychus arator) (Mattheissen and Learmonth 1995)

leaf blister sawfly (Phylacteophaga froggatti) (Farrell and New 1980) spring beetle

(Liparetrus jenkinsi and Heteronyx elongatus) wingless grasshopper (Phaulacridium

vittatum) (Loch and Floyd 2001) eucalyptus weevil (Gonipterus scuttelatus) (Took

1955) chrysomelid beetles (Paropsis spp Paropsisterna spp and Cadmus spp) (Loch

and Floyd 2001) autumn gum moth (Mnesampela privata) (McQuillan 1985)

phoracantha beetles (Phoracantha solida) (Lawson et al 2002) and psyllids

(Ctenarytaina eucalypti) (Elliot and de Little 1985)

Symptoms of damage may vary greatly between pest species For example African

black beetles cause damage to stems by removing bark just below ground level (Abbot

1993 Mattheissen and Learmonth 1995) Leaf blister sawflies cause damage to foliage

by feeding on the mesophyll tissue that occurs between the upper and lower epidermis

29

which creates a blister on the leaf lamina (Farrell and New 1980) Chrysomelid beetles

tend to scalp the edges of leaves in semi circles (Loch and Floyd 2001) and gum leaf

skeletonisers (Uraba lugens) only feed on the tissues between the leaf veins (McQuillan

1985 Farr 2002)

Compared with E globulus E dunnii has only recently been utilised as a plantation

species in southern Queensland (Jovanovich et al 2000) Since 1999 E dunnii

plantations have been increasingly impacted upon by insect pests for which there is a

paucity of knowledge (Lee et al 2000) One record by Carnegie and Angel (2005)

reported high levels of damage by Creiis lituratus (Psyllidae) in a young E dunnii

plantation in northern NSW This damage was severe enough to render much of the

plantation unfeasible to harvest for profit Phoracantha beetles and cossid moths have

also been observed causing severe damage to E dunnii plantations in southern

Queensland (Lawson et al 2002) Because the Queensland plantation industry is in its

infancy accurate identification of pests is becoming increasingly important for industry

development

Chapter Aim

The aim of the present study was to increase the knowledge of insect pests of eucalypt

plantations in southern Queensland Consequently an inventory of pest species of E

dunnii and to a lesser extent E grandis was conducted over a three-year period (2003-

2006) Profiles and general ecological information for important pest groups are

presented

30

Materials and Methods

Site Selection

Opportunistic sampling was conducted in 25 plantations over a three-year period These

plantations occurred in a range of localities along the Queensland coast from Brisbane

to Bundaberg Some minor sampling was also conducted near Casino in northern New

South Wales

Most plantations exclusively contained E dunnii however a few plantations contained

a mixture of both E dunnii and E grandis Plantations ranged from one to six years old

Other eucalypt species which were sampled to a lesser extent included E globulus E

tereticornis E tessilaris and hybrids (E grandis x camaldulensis E grandis x

tereticornis E tereticornis x urophylla E urophylla x camaldulensis and E urophylla x

grandis)

Sampling Regime

Sampling occurred during field surveys which took place every three-months from

December 2003 to November 2006 Each field survey occurred over a three-week

period Insect specimens were collected opportunistically as they were encountered in

plantations during drive-through surveys (Speight and Wylie 2001) Information was

collected daily in the field including the number and frequency of species encounters

the severity of associated damage and the state of the host Photographs were taken of

relevant species and their associated damage

Specimen Collection and Storage

Insect specimens were collected by hand which involved picking a number of

individuals from the surfaces of foliage and stems and placing them in plastic vials

31

containing 70 ethanol as a preservative Boring insects were removed from their

galleries after the stems were cut using a machete Most hard-bodied insects were

euthanized using an ethyl-acetate solution or by being placed in a freezer Some of the

larger insects were gutted pinned dried and stored in entomology boxes with

naphthalene crystals for preservation Soft bodied specimens such as larvae were

stored in 70 ethanol

Insect Identification

Insect specimens were examined at high magnification using an Olympus stereo

microscope (Olympus digitalcopy) Specimens were identified to the lowest possible

taxonomic level (in most cases to genus and species level) As the number of

specimens increased species were identified using various resources such as

entomological literature (family level Waterhouse 1970) websites (Pest and Disease

Image Library PADIL wwwpadilgovau wwwcsiroauorgentomology) with help of

taxonomic experts (S Lawson Queensland Department of Primary Industries C Reid

The Australian Museum Sydney) and by comparing specimens with voucher specimens

at the Australian Museum in Sydney

32

Results

During the survey 46 insect species were identified These consisted of 36 pest species

and ten predatory species (beneficial) Species identified as incidentals or lsquotouristsrsquo

were not included in the study Pests included 18 defoliators 13 sap-suckers three

borers one leaf blistering species and one gall forming species Only three insect

families were identified as causing high levels of damage These were the

Chrysomelidae the Miridae and the Cossidae All other species were found at either

low or medium abundance (Table 21)

Pest (P) Beneficial (B) Order Family Species Mode of feeding Abundance

P Coleoptera Cerambycidae Phoracantha solida Borer Medium

P

P

Coleoptera Curculionidae Gonipterus scuttelatus

Oxyops sp

Defoliator

Defoliator

Medium Low

P

P

P

P

P

P

P

P

Coleoptera Chrysomelidae Paropsis atomaria

Paropsis obsoleta

Paropsis variolosa

Paropsisterna cloelia

Paropsisterna agricola Longitarsus sp

Paropsisterna sp

Cryptocephalus sp

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

High

Low

Low

High

Low

Low

Low

Low

P Lepidoptera Cossidae Endoxyla cinerea Borer High

P Lepidoptera Xyloryctidae Uzucha humeralis Borer Low

P

P

Lepidoptera Limacodidae Doratifera quadriguttata Doratifera vulnerans

Defoliator

Defoliator

Low

Low

P Lepidoptera Psychidae Hyalarcta sp Defoliator Low

P Lepidoptera Tortricidae Stepsicrates sp Defoliator Medium

P Lepidoptera Geometridae Pholodes sinistraria Defoliator Low

P Lepidoptera Nolidae Gastrophora sp Defoliator Low

P Hymenoptera Tenthredinidae Phylacteophaga sp Leaf blister Medium

P Hymenoptera Chalcidoidea Ophelimus sp Gall former Medium

P Hemiptera Miridae Rayieria Sap-sucker High

P Hemiptera Eriococcidae Eriococcus coreaceus Sap-sucker Medium

P Hemiptera Pentatomidae Nezra viridula Sap-sucker Low

P

P

Hemiptera Psyllidae Glycaspis sp

Cardiaspina sp

Sap-sucker

Sap-sucker

Medium

Low

P Hemiptera Eurymelidae Eurymela fenestrata Sap-sucker Low

P Hemiptera Flatidae Siphanta sp Sap-sucker Low

P

P

Hemiptera Coreidae Mictis profana

Amorbus sp

Sap-sucker

Sap-sucker

Low

Medium

P Hemiptera Scuttigeridae Scutiphora sp Sap-sucker Low

P Hemiptera Pentatomidae Poecilometis armatus Sap-sucker Low

P Hemiptera Margarodidae Pseudococcus sp Sap-sucker Low

P Hemiptera Eurybrachidae Platybrachys sp Sap-sucker Low

P Orthoptera Acrididae Goniaea vocans Defoliator Low

P Orthoptera Acrididae Atractomorpha similis Defoliator Low

Table 21 Insects identified causing damage in eucalypt plantations in southern Queensland

33


P Orthoptera Tettigoniidae Caedicia sp Defoliator Low

B Hemiptera Pentatomidae Oechalia schellenbergii Generalist Predator Low

B Hemiptera Reduviidae Pristhesancus plagipennis

Generalist Predator Low

B Coleoptera Coccinellidae Coccinella repanda Generalist Predator Low

B Mantodea Mantidae Tenodera australasiae Generalist Predator Low

B Mantodea Mantidae Ima fusca Generalist Predator Low

B Mantodea Mantidae Rhodomantis pulchella Generalist Predator Low

B Mantodea Mantidae Orthodera ministralis Generalist Predator Low

B Neuroptera Nymphidae Nymphes myrmeleonoides


B Neuroptera Chrysopidae Mallada signata Generalist Predator Low

34

Longicorn Beetles

Order Coleoptera

Family Cerambycidae

Genus Phoracantha sp

Identification (Genus)

Small to large beetles 5-60 mm long (elongate sub-cylindrical and lightly pubescent)

Head hypognathous Antennae large distinct more than two-thirds as long as the body

(inserted on frontal prominences within emargination of the eye) and capable of being

directed backward parallel and above the body Palpi with terminal segments

subcylindrical or fusiform Pronotum with sharp edged lateral margins Fore coxae

transverse or globular separated the cavities open behind Mesepimera not reaching

mid coxal cavities All tibiae with two spurs Claws simple Elytra covering abdomen and

usually patterned with cream and brown markings

Larvae Body elongate subcylindrical slightly flattened straight lightly sclerotised

length ranging from 5-220 mm Head small and dark brown lightly sclerotised

Prothorax large and yellow Mesothorax and metathorax indistinct Legs reduced

Abdominal sides with lateral swellings or ampullae concolorous with pronotum (Figure

2-1B amp C)

Life History and Biology

The biology of many Phoracantha species is poorly known Field observations of

Phoracantha solida (native to southern Queensland) suggest that adults emerge in early

spring and lay small batches of eggs (1-5) under bark crevices of eucalypt species The

resultant larvae bore into the cambium and feed on the phloem Mature larvae tunnel

into the heartwood to pupate and adults emerge through emergence holes in the stem

(Lawson et al 2002)

35

Adult beetles are attracted to stressed eucalypt hosts which may be detected by

chemicals produced by foliage and stems Adult beetles may synchronise ovipositon

with periods of maximum water stress Low bark moisture content appears to increase

the survival rate of larvae (Hanks et al 1999)

Description of Damage

Damage is caused by larvae tunnelling within the stem which causes a reduction in

wood quality Affected trees may be identified by the presence of oval shaped

emergence holes on the surface of the stem cracks and discolouration of the bark due

to the colonisation of saprophytic fungi and frass at the base of the tree (produced by

feeding larvae) (Figure 2-1A) Affected trees may also display senescent foliage if the

stem damage reduces water transport from the roots to the crown Such trees may

eventually die

Threat to Industry

Several Phoracantha species are pests of Eucalyptus species in Australia and

overseas These include P acanthocera (Abbot et al 1991) P impavida (Curry 1981)

P semipunctata P recurva (Lanfranco and Dungey 2001) and P solida (Lawson et al

2002) In Australia P semipunctata and P recurva are mainly pests of stressed trees

such as those subjected to drought conditions (Duffy 1963 Lanfranco and Dungey 2001

Lawson et al 2002)

Three Phoracantha species are important pests in southern Queensland including P

solida (two hole borer) P acanthocera (bullseye borer) and P mastersi (ringbark

longicorn) P solida is the most widespread species within the region (Elliot et al 1998

Lawson et al 2002) Larvae of P solida were observed in approximately 50 of

plantations including many that appeared to be otherwise healthy Almost all

36

infestations were minor however two severely affected E grandis plantations were

identified Both these plantations appeared to be severely moisture stressed which may

have increased their susceptibility to beetle attack

Figure 2-1 A damage to an E grandis stem by longicorn beetle larvae (Arrow indicates brown rotting tissue infected with saprophytic fungi) B amp C longicorn beetle larvae Scale bar 5 mm

37

Eucalypt Weevils

Order Coleoptera

Family Curculionidae

Species Gonipterus sp and Oxyops sp

Identification (Species - QLD form)

Small beetles 5-8 mm in length (stout) Head hypognathous produced into a rostrum

with terminal mouthparts Antennae 3-4 segmented clubbed elbowed and inserted into

the rostrum in front of the eyes Pronotum and elytra heavily sclerotised lightly

pubescent red-brown with lighter coloured irregular tubercles (Figure 2-2B amp E)

Larvae to 7 mm in length Bright yellow with a black dorsal stripe secreted slime

covering the body Head reduced legs reduced A lsquotailrsquo of faecal matter often attached

to the posterior end of the body (Figure 2-2A)

Life History and Biology of the related species Gonipterus scuttelatus

Adults and larvae feed on expanding eucalypt foliage (Took 1955 Elliot and de Little

1984) Adults emerge in autumn and delay their egg laying until the following spring

Beetles spend the winter hiding under bark or under debris on the ground In Australia

there are generally two generations per year however in warmer climates such as

parts of Chile there may be as many as four generations of beetles per year and larval

development may occur in less than 40 days (Lanfranco and Dungey 2001)

Overwintering adult weevils reappear in spring (AugustSeptember) and begin egg

laying Egg laying by adults continues until November Eggs are laid in a pod which

hatches in 3-4 weeks The first larva to emerge chews directly through the pod and leaf

surface on which the pod was laid making a hole through which all the other larvae

emerge There are four larval stages which last for a total of approximately 4-7 weeks

depending on temperature When fully developed the larvae drop to the ground and

38

pupate in the soil forming small pupal cells or chambers made from soil particles

cemented together just below the surface The pupal stage lasts approximately 8

weeks Adults emerge from the soil in January for the second generation Larvae of this

generation mature and pupate in late summer with adult weevils emerging in

MarchApril (Phillips 1992a)


Most damage is caused by the larvae which feed on one side of the leaf by creating

irregular holes or lsquotrack marksrsquo in the leaf Adults feed on the leaf edges and give foliage

a ragged appearance (Phillips 1992) (Figure 2-2D)

Threat to Industry

The eucalypt weevil has a wide host range in the genus Eucalyptus (Took 1955 Elliot

and de Little 1984 Phillips 1992) The species is a more important pest of overseas

plantations than Australian (Took 1955 Richardson and Meakins 1986) Recent studies

show that G scuttelatus is in fact a species complex (Loch 2006) and specimens from

Queensland may represent a different species Gonipterus and Oxyops are commonly

misidentified due to their similarity (S Lawson pers comm) Given that some

individuals occurring in Southern Queensland plantations appeared to be different to

both species these may be an undescribed species (Figure 2-2E)

No serious defoliation events were observed in Southern Queensland The common

occurrence of individuals and their damage in plantations may suggest that numbers will

increase in the future This pest should therefore be monitored so that action can be

taken quickly if an outbreak does occur

39

Figure 2-2 A larva of a curculionid species feeding on Eucalyptus (the arrow indicates the faecal lsquotailrsquo) B C amp E adult weevils (arrow) D damage caused by adult beetles Scale bar 5 mm

40

Chrysomelid Beetles

Order Coleoptera

Family Chrysomelidae

Subfamilies Paropsinae Crytocephalinae

Species Paropsis atomaria Paropsis obsolete Paropsis variolosa Chrysoptharta cloelia Chrysoptharta agricola Longitarsus spp Paropsisterna spp Cryptocephalus spp

Identification (Select Species)

Small to moderate sized beetles 15-22 mm length (oval convex elongate or

cylindrical) Usually robust usually not pubescent often brightly coloured spotted or

metallic) Head prognathous or hypognathous sometimes reflexed beneath antennae

filiform moniliform slightly serrate or thickened apically without a pronounced club (not

extending past middle of the body not inserted in emarginations of the eye or on

prominences) Pronotum with or without defined margins Fore coxae globular or

transverse projecting or not closed or open behind Mesepimera usually reaching mid

coxal cavities Pair of tibial spurs not present on all legs Hind femora sometimes

dentate behind Elytra usually covering abdomen rarely exposing part of the pygidium

Wings usually present (Britton 1970)

Beetles in the Paropsinae subfamily are often recognised by their bright colours and

tortoise shaped bodies (subfamily Paropsinae) Colour patterns vary between species

(Figure 2-3A-K) The body of adults of Paropsis atomaria may vary in colour from

orange to pinkish with yellow speckles (Figure 2-3A) Paropsis obsolete has orange

elytra with purple speckled bands (Figure 2-3E) Chrysoptharta cloelia can be entirely

black orange or black with orange shoulders (Figure 2-3B) (Matthews and Reid 2002)

Adult beetles in the Cryptocephalinae have more cylindrical lsquocigar shapedrsquo bodies than

the Paropsinae (Figure 2-3C amp I) (Matthews and Reid 2002)

The larvae of the Chrysomelidae vary greatly in pattern and colour between species and

41

at different stages of development The body is usually short and sub-cylindrical with a

strongly schlerotised head capsule and three short thoracic legs (Figure 2-4C-I)

(Matthews and Reid 2002)

Biology and Ecology

Chrysomelids are pests of a large number of plant species including eucalypts (de Little

1989 Simmul and de Little 1999) Adult beetles may overwinter under bark or amongst

leaf litter In response to warmer spring temperatures the adults become active begin

feeding on foliage and begin mating Eggs are laid on the foliage or stem of the food

plant Eggs vary in shape and colour between species and may vary in the way they are

oviposted on the leaf such as in rows (Figure 2-4A) or in a rosette around the stem

(Figure 2-4B) The larvae of many species will aggregate whilst feeding on foliage

Mature larvae burrow into the soil where they pupate Adult beetles are strong fliers and

are believed to be capable of travelling several km between plantations (Matthews and

Reid 2002 Reid 2006)

Symptoms and Damage

Leaves are chewed from the margin inward towards the midrib in a semicircle (scalping)

(Figure 2-4D E F amp G) First instar larvae often feed in rows and devour entire leaves

whereas mature larvae cause damage that is similar to adult damage

Threat to Industry

Many chrysomelid species are considered important pests of Australian eucalypt

plantations (especially those in the genera Paropsisterna and Paropsis) (Tanton and

Khan 1978 de Little and Madden 1975 Loch and Floyd 2001 Nahrung et al 2001

Ohmart and Edwards 2001 Nahrung 2006) Damage caused by chrysomelid beetles

can severely affect the growth rate height volume and quality of plantation trees

42

(Candy et al 1992 Elek 1997 Elliot et al 1998) Some of the more important pests of

plantations include Paropsisterna maculata and Paropsisterna agricola which are

serious pests of E nitens plantations in Tasmania (de Little 1989) Paropsisterna

variicollis and Paropsisterna nobilitata are serious pests of E globulus plantations in

Western Australia (Selman 1994 Simmul and de Little 1999 Loch 2005) Severe

damage by Paropsisterna cloelia has been reported in E grandis plantations in coastal

New South Wales (Carne et al 1974) however Wylie and Peters (1993) did not list any

chrysomelid species causing damage in eucalypt plantations in southern Queensland in

1993 Lawson and Ivory (2000) found several emergent plantation pest species in

southern Queensland in 2000 which suggests that species have built up numbers over

the past decade Nahrung (2006) recorded 17 species of chrysomelid beetles

associated with E cloeziana and E dunnii plantations in southern Queensland The

most abundant of these species were Paropsis atomaria Paropsis charybdis and


43

Figure 2-3 A adult Paropsis atomaria B adult Paropsisterna cloelia C mating adults of Cryptocephalus sp D Paropsisterna sp (possibly a brown colour form of P cloelia) E adult Paropsis obsoleta F adult Paropsisterna sp G Altica sp) H Paropsisterna sp I adult Cryptocephalus sp J unknown species K adult Trachymela sp scale bar 5 mm

44

Figure 2-4 A eggs of Paropsisterna cloelia B eggs of Paropsis atomaria C larva of an unknown chrysomelid species D larva of Paropsisterna cloelia E larva of an unknown chrysomelid species F amp G larvae of Paropsis atomaria at different stages of development H larva of Paropsisterna sp I larva of Paropsis sp (F G amp H arrow points to typical scalping damage) All pictures scale bar 5 mm

45

Giant Wood Moths

Order Lepidoptera

Family Cossidae

Species Endoxyla cinerea

Identification (Species)

Australiarsquos largest moth species up to 20 cm length Head ocelli not present antennae

bipectinate (only in the basal half of the male) maxillary palpi minute labial palpi short

epiphysis present Thorax and abdomen covered in dense grey brown hair Wings

large strong and narrow Forewing with strong median vein forked in distal cell CuP

present Hindwing with median vein forked (Common 1970 1990)

Larva up to 15 cm length subcylindrical Prothorax with large schlerotised shield and 3

prespiracular setae Abdomen yellow or pink with transverse yellow banding (Figure

2-5D amp E) Pupa long cylindrical abdomen spined segments 3-7 movable in male 3-6

in female cremaster absent protruded from tunnel at ecdysis (Common 1970 1990)


Adult moths are active during summer months and lay their eggs on the bark of tree

stems under a glutinous secretion As many as twenty thousand eggs can be laid by a

single female (Common 1970 1990) First instar larvae produce a silken thread from

the abdomen to assist aerial dispersal It is unknown whether larvae burrow into the

ground first to feed on host roots or whether they bore directly into the host stem

Observations suggest the former as larvae within stems always appear to be more than

10 mm in length Larvae bore into the stems of trees usually aged 12 months A

chamber is excavated within the cambium which connects to a vertical tube in the

heartwood It is likely that the vertical tube is used for protection while the cambial

chamber is used for feeding (Zalucki et al 2002) A well-developed gallery usually has a

46

lsquoJrsquo shape cross-section (Figure 2-5B) Larvae feed within the gallery for 1-2 years before

pupating and emerging as adult moths (Monteith 1991b) Moths are active at night but

do not appear to be attracted to lights They may be found during the day resting on the

bark of trees (Monteith 1991a)


The most obvious indication of giant wood moth damage is the occurrence of an

emergence hole at the base of the stem (holes occur higher on the stem as the tree

grows taller) Such holes can be difficult to locate in rough barked species Larvae are

usually well developed before damage becomes conspicuous (Fig 25 A amp D) Frass

(granulated wood and waste) often accumulates at the base of the tree (Monteith

1991b) Larvae may remove a large quantity of heartwood which may weaken the stem

and may cause breakage during windy conditions (Figure 2-5C)

Secondary damage may occur due to attack by yellow tailed black cockatoos The birds

can cause severe damage by tearing into the stem with their beaks in search of the

larvae (McInnes and Carne 1978) Fungal staining and stem rot are often associated

with larval galleries

Threat to Industry

E cinerea is a serious pest of eucalypt plantations in Queensland and New South

Wales (Wylie and Peters 1993 Lawson et al 2003) Some eucalypt species are

particularly susceptible to attack including E grandis E dunnii and E grandis x E

camaldulensis (McInnes and Carne 1978 Lawson et al 2003) There is potential for E

cinerea to become an increasingly widespread pest in southern Queensland

47

Figure 2-5 A borer gallery occupied by a wood moth larva B typical lsquoJrsquo shaped gallery with a large emergence hole opening to the right of the stem C broken stem that has snapped during windy conditions due to damage caused by a wood moth larva (arrow) D large larva of Endoxyla cinerea (yellow form) E a small larva of Endoxyla cinerea (pink form) Scale bar 5 cm

48

Case Moths

Order Lepidoptera

Family Xyloryctidae

Species Uzucha humeralis

Identification (Family)

Medium sized stout moths (generally drab) Head small smooth scaled ocelli absent

antennae in male simple ciliated scape without pectin Maxillary palpi 3 segmented

Forewing with CuA2 arising well before the lower angle of distal cell Forewing pale grey

often with dark spot midway along the wing Hindwing broader than forewing black

fading to pale yellow at the margins Hind tibia with long slender scales Abdomen with

dorsal spines and often with alternating bands of black and orange

Larva up to 40 mm length sub-cylindrical Head dark brown strongly schlerotised

Prothorax paler brown than abdomen Abdomen brown to black sparsely pubescent

crotchets in elipse (Figure 2-6B) (Common 1970 Common 1990)


Eggs are laid on the bark of the host Larvae bore into the stem for a short distance (1-2

cm) to create a space used for protection against predators A silk sheet is spun around

the stem which is covered with frass to conceal the bore entrance Larvae feed mainly

on the bark (Figure 2-6A)


Damage may be recognised by the presence of a silk sheet on the surface of the stem

Discolouration of the stem is caused by the removal of bark Larvae may be detected by

removing the silk sheet from the stem

49

Threat to Industry

Damage caused by a single larva of U humeralis is generally low however large

numbers have the potential to cause severe damage Case moth larvae were often

found in plantations in both Queensland and New South Wales Individuals were rarely

collected in E dunnii plantations and were more common on E tessilaris (a common

ornamental species near homesteads)

U humeralis is currently regarded as a minor pest in southern Queensland

Figure 2-6 A damage by Uzucha humeralis a silk and frass tent is constructed (arrow) and brown discoloured bark occurs above B larva of Uzucha humeralis (Scale 5 mm)

50

Cup Moths

Order Lepidoptera

Family Limacodidae

Species Doratifera quadriguttata amp Doratifera vulnerans


Small stout moths 10-20 mm length Head ocelli and chaetosemata absent antennae

bipectinate in male haustellum small maxillary palpi 1-3 segmented labial palpi short

and 2-3 segmented epiphysis absent Thorax and abdomen densely pubescent

Forewing broad with M present in discal cell forked chorda absent R3 R4 and R5

stalked Hindwing with M present in discal cell rarely forked Sc and R1 fused with Rs

near base or connected to Rs by R1 CuP present (Hadlington 1966 Common 1970

Common 1990)

Larvae to 15 mm length squat patterned and colourful subcylindrical Head

hypognathous and retracted antennae long Thoracic legs reduced prolegs absent

ventral suckers on abdominal segments 1-7 Setae modified and forming stinging hairs

dorsally in groups at posterior and anterior end of the body

Doratifera quadriguttata is leaf green with a row of intricate black and white markings

surrounded by stinging hairs on dorsal ridges (Figure 2-7C amp D) Hairs produce rash like

symptoms if brushed against bare skin

Doratifera vulnerans has two large brown patches at either end of the body and a

central patch of bright yellow Six clusters of red stinging hairs occur at either end of the

body on raised ridges (Figure 2-7E) (Hadlington 1966 Common 1970 Common 1990)


Limacodid moths are often called cup moths because of their characteristic cup shaped

pupal case (Figure 2-7A amp B) Adult moths are active at night and are readily attracted

51

to lights Eggs are laid on the host and the resultant larvae feed on foliage (especially

flush growth) When the larvae pupate they spin a hard smooth pupal case resembling

a eucalypt fruit (potentially mimicry) Moths emerge from pupae through an operculum

and the remaining case resembles a cup (Figure 2-7D) (Hadlington 1966)

Symptoms and Damage

Immature larvae skeletonise foliage by feeding on one side of the leaf and avoiding

veins and the midrib Mature larvae feed on the whole leaf lamina including the midrib

Threat

Although cup moths can cause severe defoliation to trees in rural areas there is very

little information about impacts in eucalypt plantations (Hadlington 1966 Southcott

1978 Ohmart and Edwards 1991) Pook et al (1998) observed a severe outbreak of

larvae in Corymbia maculata forest in southern New South Wales

At least four species of cup moths were observed in plantations in southern

Queensland The most abundant species were Doratifera quadriguttata (Figure 2-7A C

amp D) and Doratifera vulnerans (Figure 2-7B) These species were encountered

frequently in plantations but did not occur in great numbers Cup moths are considered

minor pests

52

Figure 2-7 A developing pupa of Doratifera sp B an emerged pupal case of Doratifera sp (arrow indicates emergence hole) C D larva of Doratifera quadriguttata E larva of Doratifera vulnerans (arrows indicates stinging hairs) Scale bar 5mm

53

Leaf Bag Worms

Order Lepidoptera

Family Psychidae

Species Hyalarcta huebneri

Identification of Species

Medium sized stout moths 10-20 mm length Female is apterous without legs or

developed antennae occupying pupal case as an adult Male is mobile and capable of

flight Head covered in dense rough hair and blackish ocelli large tongue obsolete

antennae strongly bipectinated to apex and dark orange labial palpi very short Thorax

covered in dense rough hair and blackish posterior tibiae without middle spurs (end

spur is short) Forewings elongate triangular costa straight apex rounded vein 1a

anastomosing with vein 1b before middle vein 1c coincident with vein 1b beyond

middle vein 5 absent vein 6 from above middle transverse vein vein 7 sometimes out

of 8 veins 8 and 9 present stalked veins 10 and 11 sometimes stalked Hindwings

small termen rounded costa with a broad black line from base to middle vein 5 absent

veins 6 and 7 approximated or coincident 8 coincident with one costal pseudoneuria

(Meyrick and Lower 1907 Common 1970 Common 1990)

Case length 40-45 mm diameter 13-15 mm circumference 30 mm ovate lanceolate

broadly tapering at both ends dull grey whitish or dark grey thickly ornamented apart

from the posterior 16 Ornamented with large pieces of eucalypt leaves placed

indiscriminately (Figure 2-8) Opening broadly ovate fixed to the food plant by strong

silk (Meyrick and Lower 1907 Common 1970 Common 1990)

Larva dark grey black second thoracic segment irrorated with dark grey conspicuous

red anal segments (Meyrick and Lower 1907 Common 1970 Common 1990)

54

Life History and Ecology

The larvae of bagworms construct a case from plant material such as leaves and twigs

which are spun with strong silk which is secreted by the larva (Figure 2-8A) Case

material is collected from the host plant and is often characteristic of the species The

case serves to camouflage the larva from predators and new material is added to the

anterior end of the case as the larva grows Larvae are mobile within the case and may

move by pushing the anterior end of the body through an opening at the end of the case

and dragging it around This allows the larva to feed on its host plant while remaining

protected Female larvae pupate and live within the case their entire lives and are

wingless Male moths emerge and are the dispersing sex (Heather 1975)


Leaf bagworms tend to feed on the entire leaf including the midrib Larvae may partially

eat a leaf before moving onto another Severely damaged canopies may acquire a

ragged appearance due to bagworm defoliation

Threat to Industry

Only two species occur within the genus Hyalarcta (Nielsen et al 1996) These species

are known to feed on over 40 species of Australian plants (Heather 1975 1976) H

huebneri has a large distribution from north-east Queensland to eastern New South

Wales Victoria south-east South Australia and south-west Western Australia H

huebneri has caused severe damage in Pinus radiata plantations (Heather 1975 1976)

but here are no records of damage in eucalypt plantations

On one occasion H huebneri was found causing high levels of damage in southern

Queensland In early summer a small group of trees in a two-year-old plantation of E

grandis were almost completely defoliated (95 defoliation) This was the only instance

55

in which the species was encountered however the large feeding capacity of this

species indicates that it has the potential to become an important pest in plantations

Figure 2-8 A pupal cases of Hyalarcta hueberli ornamented indiscriminately with eucalypt leaves (arrows indicate the end from which the larvae partially emerge to feed) Scale bar 5 mm

56

Eucalypt Leafrollers

Order Lepidoptera

Family Tortricidae

Species Stepsicrates sp


Slender delicate day flying moths 10-25 mm length Head rough scaled ocelli present

antennae clubbed chaetosemata present maxillary palpi 2-4 segmented labial palpi

short to very long rarely ascending apical segment short and obtuse Thorax black

Abdomen black with orange bands Forewing black with patches of white near the

apex with costa strongly arched costal fold present in male chorda and M present

CuA2 arising before three fourths of distal cell Cup near margin Hindwing pale brown

with pectin of hairs on CuA CuP present

Larva elongate cylindrical to 25 mm Head capsule dark brown and strongly sclerotised

Thoracic legs and abdominal prolegs present crochets uniordinal Abdomen with

alternating dark and light brown longitudinal bands (Figure 2-9B amp C)

Pupa with spined abdomen cremaster with hooked spines (Common 1970 1990)


Adult moths are active during the warmer months during daylight Stepsicrates species

are commonly called magpie moths due to the black and white markings on their wings

Eggs are scale like and laid individually on stems or foliage The first instar larvae are

cryptic and produce strong silk to bind leaves thereby creating a refuge These leaves

accumulate as the larva develops and may contain both living and dead tissue The

larvae feed on foliage from within the refuge

57


The damage caused by leafroller caterpillars is very distinctive The leaves at the end of

branches are bound together to form an aggregation of foliage The larva produces

copious faecal pellets which also stick to the silk (Figure 2-9A) Both the binding of

leaves and defoliation by the larva is likely to have an adverse effect on host growth

Threat to Industry

There are no records of severe damage by Leafroller caterpillars in Australian eucalypt

plantations however Strepsicrates macropetana is an Australian species which has

been introduced to New Zealand where it causes high levels of damage in young

plantations (Philpott 1923 Nuttall 1983 Mauchline et al 1999)

Leafroller caterpillars were abundant in younger plantations in southern Queensland It

was observed that single larvae could cause surprisingly severe levels of damage

Larvae appeared to prefer new growth in one and two-year-old plantations Incidence

appeared to be greater in the northern plantations where the climate is warmer

Strepsicrates sp appears to have the potential to become an important pest in southern

Queensland

58

Figure 2-9 A an aggregation of eucalypt foliage spun with silk from Strepsicrates sp B Strepsicrates sp larva spinning silk to bind a leaf which occurs at the far right of the picture (arrow points to silken threads) C mature larva Scale bar 5 mm

59

Leaf Blister Sawfly

Order Hymenoptera

Family Tenthredinidae

Species Phylacteophaga sp


Small wasps 4-8 mm length Head without subantennal grooves Tergum 1 distinct

though closely associated with mesanotum prepectus defined Cenchri protruding from

mesanotum Mesoscuttellum not separated from scutum laterally and axillae not defined

anteriorly Forewing with a second incomplete anal cell Hindwing with basal field

strongly developed and emarginate Nygmata present Abdomen broadly sessile at

base no marked distinction between segments 1 and 2 (Riek 1970)

Larva stout dorso-ventrally compressed Head dark Thorax grey with short black

thoracic legs Crochets absent Abdomen pale grey sparsely pubescent (Riek 1970)


Leaf blister sawflies are active during the warmer months and may produce several

generations a year Adults live for less than a week and do not feed Eggs are laid

singly on foliage of the host The resultant larvae chew through the cuticle into the inner

leaf tissues and develop within a leaf cavity beneath the cuticle As the cavity increases

in size it takes on the appearance of a blister Mature larvae pupate within the cavity

and emerge through a hole in the cuticle (Farrell and New 1980 Thumlert and Austin

1984)


Blisters caused by leaf blister sawflies are easily recognisable and may be up to 10mm

in diameter (Figure 2-10A B amp C) A small lsquotailrsquo on one side of the blister often occurs

where the cavity was initiated by the first instar larva (the lsquotailrsquo is diagnostic of

60

Phylacteophaga eucalypti)

Threat to Industry

Leaf blister sawfly is a pest of E globulus plantations in the eastern states but will

usually cause only cosmetic impacts Damage mainly occurs on lower branches which

has less effect on tree growth (Loch and Floyd 2001)

Leaf blister sawflies were uncommon in southern Queensland and infestations generally

affected only a few trees in older plantations (4-5 years) The species is currently

regarded as a minor pest

Figure 2-10 A B C damage caused by leaf blister sawfly Phylacteophaga sp (arrows indicate exit holes of adult sawfly) Scale bar 5 mm

61

Eucalypt Gall Wasps

Order Hymenoptera

Superfamily Eulophidae

Species Ophelimus sp


Small wasps 1-3 mm length (dark iridescent green with transparent wings) Pronotum

with large degree of movement with the mesothorax prepectus large and slightly

convex caudally with impressed margins antennae thickened throughout flagellum with

a distinct club Femora with an enlarged bristle at the apex Fore tibial spur short and

straight basitarsus with a strigil modified into an oblique comb at the base mid tibial

spur enlarged tarsi 4 segmented (Riek 1970)

Larvae 1-4 mm length pale cream to white with distinct segmentation Reduced head

and mouthparts Legs absent


Eucalypt gall wasps are parasitic wasps and inject their eggs into foliage and stems

Trees respond by creating a malformation of tissue around the egg forming a gall

(Figure 2-11A amp B) Within the gall resultant larvae feed on the inner tissues Larvae

pupate and emerge through a hole in the gall surface (Figure 2-11B)

Other wasps may parasitise gall wasps and it is therefore important to recognise that

causal species and secondary species may be confused


Wasp galls are easily recognisable and consist simply of a spherical malformation on

the leaf or stem of the host plant Some galls remain the same colour of the leaf while

others are powdery and pinkish in colour The presence of emergence holes on the

62

surface of the gall indicates that wasps have emerged

Threat to Industry

Galls may be caused by a large number of wasp species Approximately 50 species of

Ophelimus are pests of eucalypts (Withers et al 2000) E globulus is the most

susceptible plantation species to gall wasps (Withers et al 2000) At least five species

of gall wasps are important pests of eucalypts occurring outside Australia (Flock 1957

Timberlake 1957 Huber et al 2006)

Wasp galls were frequently encountered in plantations is southern Queensland and

were usually found in younger plantations where they generally affected new expanding

foliage Wasp galls are considered to be a minor pest in plantations in southern

Queensland

Figure 2-11 A an aggregation of pink wasp galls B a green wasp gall caused by Ophelimus sp with an emergence hole (arrow) both of these are likely to be the same species Scale bar 5 mm

63

Mirid Bugs

Order Hemiptera

Family Miridae

Species Rayieria sp

Identification ( Species)

Small bugs 6-7 mm length (slender delicate) Head hypognathous black ocelli absent

Antennae filiform Maxillary and mandibular stylets elongate Thorax orangered Legs

black slender tarsi brown and 3 segmented Wings darkly tinted and transparent

hemelytron with cuneus membrane of hemelytron with 1 closed cell near cuneus

Abdomen mostly black with lateral white spots (Figure 2-12A amp C)

Nymphs resemble adults but are paler and lack fully developed wings


Mirids are active in summer and may produce multiple generations during the warmer

months Eggs are presumably either oviposited on or injected into foliage (Woodward et

al 1970) All stages are sap-sucking Adults resemble brachonid wasps and may be

mimics

Symptoms and Damage

Sap-sucking by mirids may cause vein limited necrosis of the leaf The leaf surface

becomes speckled with necrotic patches which may eventually coalesce into larger

patches (Figure 2-12B)

Threat to Industry

Several species cause damage to agricultural plants (Eyles 1999) Helopeltis spp feed

on many forest species including eucalypts and are widely distributed throughout Asia

and the pacific (Griffiths et al 2004) There are currently no records of Rayieria spp

64

causing damage in Australian eucalypt plantations

Large numbers of Mirids (Rayieria sp) were observed causing high levels of damage in

plantations in southern Queensland The damage mainly occurred in northern

plantations while plantations occurring farther south received less damage This may

be attributed to higher temperatures and rainfall in the north Mirids appear to be prolific

and should be regarded as an important pest of plantations

Figure 2-12 A a mating pair of Rayieria sp B necrotic speckling caused by Rayieria sp (arrows) C a side profile of an adult Rayieria sp Scale bar 5 mm

65

Brown Scale Insects

Order Hemiptera

Family Eriococcidae

Species Eriococcus coriaceus


Small bugs 2-4 mm length (globular and enclosed in a felted sac) Colours vary from

yellow to dark brown and red when fully mature (Figure 2-13A) The capsule of the male

is brown and waxier (Figure 2-13A) than that of the female which is white and more

cottony (Figure 2-13C) Body membranous slightly oval and elongated at the anal end

anal lobes prominent and slightly sclerotised Antennae seven segmented apex slightly

swollen legs moderately long and well developed inner margin of claw with a fine pair

of digitules longer than the claw and a small denticle near the tip (Patel 1971)


First instar nymphs emerge from eggs within the capsule of the parent female The

nymphs then travel a short distance along the branch before attaching to feed and begin

excreting their own capsule As the nymphs develop they shed their capsules and

migrate further along the stem to produce larger capsules (Woodward et al 1970 Patel

1971)

Different sexes often form separate colonies on the host At adulthood the females

remain within their capsule while the males are winged and more mobile The males

migrate between hosts to find female mates (Woodward et al 1970)

Scale insects excrete excess sugar as they feed and the droplets often called

honeydew may be collected by other insects such as ants The ants reciprocate by

providing protection from predators (Eastwood 2004)

66


Colonies of scale insects are easily recognised as aggregates of brown or white

capsules on the surfaces of stems or foliage Black sooty mould (Saprophytic

Ascomycetes) often occurs on honeydew which sticks to leaves and branches near

colonies (Figure 2-13B)

Threat to Industry

Scale insects are common pests of eucalypt plantations in Australia and have been

introduced to New Zealand (Loch and Floyd 2001 Withers 2001) Carne et al (1974)

reported severe localised damage in E grandis plantations in northern New South

Wales

E coriaceus was very common in plantations in southern Queensland Unlike foliar

pests the damage caused by scale insects is internal and cannot be visually assessed

However considering the high density of colonies it is likely that E coriaceus was

negatively affecting the growth of some trees Studies show that although evidence of

damage by scale insects is not conspicuous on foliage and stems root growth of the

host may be significantly reduced (Vranjic and Gullan 1990) E coreaceus could

become a more widespread pest in plantations in southern Queensland and it should

therefore be considered to be a potentially important pest

67

Figure 2-13 A brown scale insects Eriococcus coriaceus associated with a stem (males) B brown scale insects E coriaceus associated with a leaf midrib (sooty mould has also become associated) C brown scale insects E coriaceus (females) tended by ants (red arrow) Scale bar 5 mm

68

Green Vegetable Bugs

Order Hemiptera

Family Pentatomidae

Species Nezara viridula


Moderate to large bugs 10-15 mm length (stout) Antennae exposed from above and

five segmented ocelli well separated labium with basal segment straight Pronotum

with slender anterior projections extending to near eyes not covering scutellum

scutellum triangular and reaching apex of clavus mesosternum without median carina

hemelytra without cuneus Hindwings without hamus tarsi three segmented Generally

ime green in colour (Figure 2-14A amp B) but less frequently pale brown (Figure 2-14C)


Adults overwinter during the colder months and become active during spring when they

begin searching for food and potential mates (Drake 1920) Females have been

observed travelling over 1000m a day in search of food and oviposition sites (Kiritani

and Sasaba 1969) Eggs are yellow and are oviposited on the host in dense polygonal

clusters Incubation may be as short as 5 days in warm conditions (Harris and Todd

1980) First instar emergent nymphs aggregate near the egg mass and do not feed until

the first moult after which they disperse in search of food The nymphs feed by sap-

sucking and go through five instars before reaching adulthood which may occur in as

little as 35 days under optimum conditions (Drake 1920)


Sap-sucking causes necrotic speckling of foliage A mosaic like pattern of necrotic

patches can occur in severe infestations

69

Threat to Industry

The green vegetable bug is a serious pest of many agricultural crops worldwide

(especially legumes) (Todd 1989 DeWitt and Armbrust 1978) The literature pertaining

to the species is vast a bibliography by DeWitt and Godfrey (1979) lists over 690

references

Crops occurring near plantations in southern Queensland are known to be affected by

N viridula outbreaks during summer A single outbreak of the species was observed in

one plantation The outbreak caused low levels of localised damage and trees

recovered quickly Green vegetable bugs are not considered an important plantation

pest in southern Queensland but given that the species has the potential to be a serious

pest of a wide range of species its occurrence in plantations should be monitored

closely

Figure 2-14 A amp B adults of Nezara viridula associated with foliage (green colour form) C an adult Nezara viridula (brown colour form) Scale bar 5 mm

70

Psyllids

Order Hemiptera

Family Psyllidae

Species Glycaspis sp amp Cardiaspina sp


Small bugs 1-2 mm length (dorso-ventrally compressed) Head with broad set eyes and

a distinct median suture Elongate mouthparts forming piercing stylets Antennae 10

segmented Wings membranous hind wings with evident veins but no closed cells (M

and Cu forked) clavus present Tarsi 2 segmented

Nymphs produce a lerp a protective covering attached to the leaf beneath which the

nymphs feed The lerp is constructed from a secretion produced at the terminal end of

the abdomen The secretion consists mostly of excess sugar collected by the nymph

during feeding Different species construct characteristic lerps which aid identification

Lerps of some Glycaspis species are dome shaped with rough walls (Figure 2-15A amp

C) The lerps of some Cardiaspina species are intricately woven and basket-like (Clark

1962) (Figure 2-15C amp D)


Female psyllids lay clusters of stalk shaped eggs on foliage The resultant nymphs

move around the foliage to find suitable feed sites and begin sap-sucking and construct

a lerp The lerp increases in size as the nymphs go through five instars before reaching

adulthood Only the nymphs of psyllids produce lerps and the adults are winged and

disperse between hosts to mate and lay eggs (Woodward et al 1970)


The lerps of psyllids are distinct due to their shiny appearance At high densities

feeding psyllids can cause leaves to produce anthocyanins which causes foliage to

71

become red This is often perceived as a symptom of host stress (Sharma and Crowden

1974)

Threat to Industry

Several psyllid species cause high levels of damage in Australian eucalypt plantations

including Ctenarytaina spp Glycaspis spp Creiis spp and Cardiaspina spp (Clark

1962 Ohmart and Edwards 1991 Brenan et al 2001 Collet 2001 Yen 2002 Rao et al

2001 Carnegie and Angel 2005) Creiis lituratus has been identified causing high levels

of damage to E dunnii plantations in southern Queensland and northern New South

Wales (Carnegie and Angel 2005) This species is most active in autumn and winter

and may occur at high densities (Carnegie and Angel 2005)

Psyllids were rarely observed in plantations in southern Queensland during the study

One outbreak caused by Glycaspis sp was observed on E grandis in northern NSW In

this instance crown damage was estimated to be approximately 80 The foliage of

these trees was red due to the production of anthocyanins

E grandis plantations appear to be less susceptible to psyllids compared with E dunnii

Psyllids are considered to be a moderately important pest in southern Queensland and

their abundance should be monitored

72

Figure 2-15 A amp B nymph of Glycaspis sp (arrow points to lerp) C amp D nymph of Cardiaspina (arrow points to lerp) Scale 5 mm

73

Leafhoppers

Order Hemiptera

Family Eurymelidae

Species Eurymela fenestrata


Medium sized bugs 12-15 mm length (stout) Head black with broad set bright red

eyes Ocelli on the ventral surface of the head Tegmen shiny black with white patches

median vein extending to apex Legs long and held tightly under the body prominent

spines on the base of the hind tibiae

Nymphs with small wing buds are more brightly coloured than adults with more orange

visible dorsally


Adult females are active during the warmer months and lay their eggs in slits in the host

stem The resultant nymphs tend to aggregate during early instars (Figure 2-16A)

Nymphs go through five instars before they reach adulthood (Woodward et al 1970)

Leafhoppers are usually attended by ants which collect the sugary honeydew secreted

by the leafhoppers through the tip of their abdomens In return the ants provide

protection from predators This is a mutually beneficial relationship (Rozario et al 1993)

(Figure 2-16B amp C)


Damage to the host occurs mainly due to egg laying and consists of small necrotic

scars on stems Damage may also result from sap-sucking especially if large numbers

of insects occur

74

Threat to Industry

E fenestrata is the most common Eurymela species found on eucalypts Although large

numbers may occur in native vegetation there are very few records of damage in

eucalypt plantations Carne et al (1974) reported high numbers of an unidentified

Eurymela species causing damage in E grandis plantations in northern New South

Wales Although leafhoppers were very common in plantations in southern Queensland

they generally occurred in low numbers (aggregates of 5-30 individuals on the lower

branches of trees) and caused low levels of damage Leafhoppers are therefore

considered to be a minor pest in southern Queensland

Figure 2-16 A first instar leaf hopper nymphs Eurymela fenestrata B C adult leaf hoppers Eurymela fenestrata tended by ants (Iridomyrmex sp) (arrow) Scale bar 5mm

75

Planthoppers

Order Hemiptera

Family Flatidae

Species Siphanta sp


Small bugs 8-10 mm length (laterally compressed) Head elongate and conical clypeus

lacking lateral carina Tegmina pale green to brown held in tent like position (Figure

2-17A B amp C) strongly cross veined in precostal area strongly bent granulate clavus

Legs second tarsus small with a single small spine (Fletcher 1985)


Adults are active in summer Females attach an egg mass to the leaf lamina of the host

plant and resultant first instar nymphs will aggregate but eventually disperse as they

develop (Woodward et al 1970) Nymphs go through five instars before reaching

adulthood and mates tend to stay together When motionless the adult plant hoppers

are inconspicuous and resemble plant structures on the stem


Damage is inconspicuous and consists of tiny scars on the stem where the mouthparts

pierce the bark during sap-sucking

Threat to Industry

There are no records of Siphanta spp causing damage in Australian eucalypt

plantations Although plant hoppers were very common in southern Queensland they

occurred at extremely low densities (2-3 individuals per tree) Siphanta sp is therefore

considered to be a minor pest

76

Figure 2-17 A B adult Siphanta sp (green form) C adult Siphanta sp (brown form) Scale bar 5 mm

77

Clown Bugs

Order Hemiptera

Family Coreidae

Species Mictis profana and Amorbus sp


Large bugs 10-25 mm length (stout robust) Head half as wide as pronotum bucculae

extending behind antennifers Antennae six segmented and inserted dorsally between

the center of the eyes Membrane of hemelytron with six longitudinal veins Metapleural

scent gland openings with distinct peretremes Femora and tibia enlarged Posterior

margins of abdominal terga 4 and 5 produced posteriorly in mid line (Figure 2-18B amp C)

Nymphs tend to be more brightly coloured than adults with orange bands on the joints of

the legs (undeveloped wings) (Figure 2-18A)


Adults overwinter and are active during the warmer months There may be one to two

generations in a year Females cement their eggs to the underside of foliage of the host

plant The resultant nymphs feed by sap-sucking on the tips of foliage and go through

five instars before reaching adulthood (Woodward et al 1970) The habit of feeding at

the tips of foliage has given these insects the name tip feeders Adults are usually found

solitary or in pairs while nymphs tend to form aggregates (5-10 individuals) Adult

males display by waving their colourful hind legs to attract potential mates (Figure

2-18B)


Feeding occurs at the tips of foliage and may cause foliar and stem necrosis The

necrosis may spread partway down the leaf lamina towards the petiole

78

Threat to Industry

M profana feeds on a range of plant species both native and introduced and has a

preference for plants in the Mimosaceae family (Flanagan 1994) The species is known

to feed on E grandis and E camaldulensis (Griffiths et al 2004) Some species of

Amorbus such as A obscuricornis feeds exclusively on eucalypts (Steinbauer et al

1998)

M profana was generally found in low abundance in plantations in southern

Queensland and is currently considered to pose a low threat Interestingly some insects

were found in association with Acacia species within the inter-rows of plantations The

plantation trees occurring near these acacias were also found to have insect damage

Figure 2-18 A nymph of Amorbus sp B C adult Mictis profana Scale bar 5 mm

79

Assassin Bugs

Order Hemiptera

Family Reduviidae

Species Pristhesancus sp


Large bugs 10-25 mm length Head elongate hypognathus with distinct gulla labium

with straight basal segment elongated maxillary and mandibular stylets Pronotum not

covering scuttellum with large lateral spines prosternum with median stridulatory

groove extending to anterior of coxae Hemelytron without cuneus with 2 large cells

occupying most of the membrane Legs slender tarsi 3 segmented Abdomen dorsally

convex (Figure 2-19A B amp C)


Assassin bugs in the genus Pristhesancus are some of the largest species in the

Reduviidae family (James 1994) Adults lay eggs on foliage and the resultant nymphs

quickly disperse and are solitary Assassin bugs feed on other insects by piercing the

exoskeleton with their curved proboscis and sucking out the body fluids (Figure 2-19C)

These insects usually move quite slowly and rely on stealth rather than speed to

capture their prey (Woodward et al 1970)

Role in Plantations

Assassin bugs have a large feeding capacity and are potential biological control agents

(James 1994 Grundy and Maelzer 2000) On numerous occasions assassin bugs were

observed feeding on larvae and adult chrysomelid beetles These appeared to be the

preferred prey of assassin bugs in plantations Such predation may benefit plantations

by reducing chrysomelid numbers

80

Figure 2-19 A B adult Pristhesancus plagipennis C adult Pristhesancus plagipennis feeding on an adult Paropsisterna cloelia (arrow) Scale bar 5 mm

81

Ladybird Beetles

Order Coleoptera

Family Coccinellidae

Species Coccinella repanda


Small beetles 5-7 mm length (oval convex) Head partially covered by anterior margin

of pronotum Antennae with an apical club apical segment of maxillary palp securiform

Prothorax with distinct lateral margins lateral edges of pronotum and elytra forming an

acute angle Elytra shiny orange-red black on outer margins and along dorsal line Fore

coxae much closer together than hind coxae mid coxal cavities closed by the

mesepimera in addition to the sterna tarsi 444 claws appendiculate (Figure 2-20A)

Larva bluish grey with rows of rounded tubercles protruding dorsally along the body

(Figure 2-20B)


Adults and larvae are predatory Eggs are oviposited on host plants which have

associated prey species such as psyllids and other small insects Some species are

effective biological control agents in the glasshouse (Hagen 1962) The efficiency of

coccinellids as predators is largely attributed to their mobility and large feeding capacity

(Obrycki and Kring 1998 Baker et al 2003)

Role in Plantations

Coccinellids were often found feeding on chrysomelid larvae in plantations in southern

Queensland Several species were observed but C repanda was the most common

species This species is beneficial in plantations where it reduces pest numbers

82

Figure 2-20 A adult of Coccinella repanda B larva of Coccinella repanda Scale bar 5 mm

83

Praying Mantids

Order Mantodea

Family Mantidae amp Amorphoscelidae

Species Tenodera australasiae Ima fusca Rhodomantis pulchella amp Orthodera ministralis


Small to large insects 5-250 mm length (elongate and delicate to squat and robust)

Head hypognathus mobile triangular with large eyes antennae slender Prothorax

narrow elongate and moveable on the mesothorax Mesothorax and metothorax

similar Tegmen narrow hind wings broad and membranous Mid and hind legs slender

and unspecialised fore legs raptorial (coxae elongate and mobile femora robust and

generally spined ventrally tibia with ventral spines and a sharp apical hook) (Key 1970

Rentz 1966)


Eggs are oviposited within excreted foam which becomes hard and forms an ootheca

The ootheca may be attached to a branch stem or any other hard surface Resultant

nymphs are predatory and may be cannibalistic They are quick to disperse upon

hatching and are solitary (Key 1970 Rentz 1996) Mantids are fast moving and use their

raptorial forelimbs to catch prey (Key 1970 Rentz 1996)

Role in Plantations

Mantids are beneficial to plantations because they feed on a range of pest species

Tenodera australasiae (Figure 2-21A) and Orthodera ministralis were the most common

species in plantations in southern Queensland

84

Figure 2-21 A an adult Tenodera australasiae B an adult Ima fusca C an adult Rhodomantis pulchella Scale bar 5mm

85

Lacewings

Order Neuroptera

Family Nymphidae amp Chrysipidae

Species Nymphes myrmeleonoides amp Mallada signata


Large to small insects 10-50 mm length (elongate with long transparent wings) Head

with large compound eyes ocelli absent antennae filiform or clubbed mouthparts

simple maxillary palp 5 segmented Prothorax freely moveable varying from transverse

to very long mesothorax and metathorax well developed Legs mostly cursorial but

raptorial in the Mantispidae family Wings variable but mostly membranous and multi-

veined (two pairs of equal length) Abdomen elongate 9-10 segmented (Figure 2-22C amp

D) (Riek 1970)

Nymphs of lacewings are usually squat with large protruding mandibles The head is

broad and the body is dorso-ventrally compressed


Adults are active during the warmer months Eggs are oviposited on foliage and stems

and are attached by long stalks (Figure 2-22A) The stalks of the eggs are believed to

reduce predation by foraging insects such as ants (Riek 1970) Nymphs are predatory

and some species are arboreal while others are subterranean (Riek 1970) Nymphs of

species commonly called lsquoantlionsrsquo burrow into loose sandy soil and create a funnel

shaped trap on the surface which unsuspecting ground dwelling insects such as ants

may fall into (Figure 2-22B) The antlions wait with open jaws beneath a layer of sand at

the base of the funnel to capture their prey Adult lacewings are also predacious and

may capture prey on the wing

86

Role in Plantations

Lacewings may benefit plantations by reducing numbers of smaller pest species such

as psyllids and leaf hoppers Several lacewing species were observed in plantations in

southern Queensland The most common species was the golden eyed lacewing

Mallada signata (Figure 2-22D)

Figure 2-22 A lacewing eggs B sand traps of antlion nymphs C an adult Nymphes myrmeleonoides D an adult Mallada signata Scale bar 5 mm

87

Discussion

Summary of Important Pests

Chrysomelid beetles caused high levels of damage in southern Queensland (especially

in younger plantations with new flush growth) The most abundant chrysomelid species

was Paropsisterna cloelia Several colour forms of this species were observed which

may indicate a species complex A better understanding of the life cycle of this species

would be required to develop efficient control methods

The main cause of foliar necrosis was caused by mirids (Rayieria sp) which were

observed causing high levels of damage in plantations in southern Queensland Mirids

were prolific in plantations and caused damage both as adults and nymphs The most

damaging borer species was Endoxyla cinerea This species caused stem damage to

many two and three-year-old plantations E cinerea appeared to have a preference for

E grandis rather than E dunnii hosts Some borer species have been efficiently

controlled using biological control agents (Eldridge et al 1995) A better understanding

of the life cycle of this species would be required to develop a control method

Although the diversity of pests in southern Queensland was high the majority of species

caused low levels of damage Collectively however damage caused by pest

assemblages may be significant (discussed in future chapters)

Chemical Control of Pests

Selecting the most suitable method of pest control depends on the species involved and

the severity of the infestation Important considerations include cost efficiency and

potential effects to the surrounding environment and non-target organisms (Elliot et al

1992 Stone 1993 Eldridge et al 1995)

88

Pests are currently controlled in southern Queensland by aerial application of the

chemical Dimethoate (Rogor reg) A number of other insecticides are also available and

widely used including alpha-cypermethrin (Fastac reg or Dominex reg) Supracide and

Nuvacron These chemicals are usually applied as aerial sprays and have a lsquoknock

down effectrsquo on pests (Eldridge et al 1995) Contact insecticides such as Malathion

require contact with the body of the target pest before taking effect on the nervous

system (Eldridge et al 1995) Some borer species may be resistant to aerial spraying

with such chemicals because they are protected within the heartwood of the stem

Controlling borers sometimes requires a stem injection of insecticide such as Azodrin

(Urquhart and Stone 1995) Brown scale insects (Eriococcus coriaceus) usually occur

lower in the canopy of affected trees where aerially applied contact insecticides are

often ineffective Systemic insecticides are most effective for controlling these insects

because they are absorbed by host tissues and affect insects as they feed Systematic

insecticides are also particularly effective against other sap-sucking species such as

psyllids (Eldridge et al 1995)

Toxins produced from the spores or endotoxins of the bacterium Bacillus thuringiensis

may be used as biological insecticides These may be applied as aerial sprays and to

control several important pests of plantations (Waterson and Urquhart 1995) One

benefit of biological insecticide is that it only targets feeding insects It is therefore less

harmful to beneficial insects wildlife and humans (Eldridge et al 1995 Waterson and

Urquhart 1995)

The timing of insecticide application is important and must coincide with the vulnerable

stages of the pest life cycle For many species insecticides should be applied during

the early stages of insect development before severe damage has been inflicted (Farrell

and New 1980) The main disadvantage of using generalist insecticides such as

89

pyrethroids is that they often kill the natural enemies of pests which can lead to greater

outbreaks in successive years (Elliott and Greener 1994)

Cultural Control of Pests

Controlling pests may also be achieved by using more creative methodologies without

the use of chemicals Cultural control usually requires a detailed knowledge of the pest

life cycle to identify vulnerability For example Phoracantha beetles are attracted to

stressed trees which exhibit senescing foliage (Duffy 1963 Lanfranco and Dungey 2001

Lawson et al 2002) The beetles can therefore be controlled by placing newly cut

eucalypt logs in piles a short distance from plantations and allowing adult beetles to

colonise and lay their eggs The logs are then burnt to destroy the insects (Soria and

Borralho 1997) Some hemipteran pests such as Eriococcus coriaceus and Eurymela

fenestrata are attended by ants which collect the lsquosugarrsquo which is excreted by the bugs

Studies show that if tending ants are controlled this can also lead to control of the bugs

(Rozario et al 1993)

Beneficial Insect Species

Pests may be controlled by insects which naturally occur in plantations These may

include predatory species such as assassin bugs and ladybird beetles or parasitic

species such as parasitoid wasps and tachinid flies Pests are also controlled by

adverse environmental conditions such as prolonged wet conditions shortage of food

and overcrowding which may result in a higher mortality rate and greater susceptibility

to diseases such as viruses (Goodyer 1985) Biological control agents may be used

which prey upon or parasitises pest species (Faulds 1991 Dahlsten et al 1998 Obrycki

and Kring 1998 Rivera et al 2001 Protasova et al 2007)

Several biological control agents have been successfully used to control pests in

90

eucalypt plantations These include bacteria (Bacillus thuringiensis) (Elliot and Greener

1994) parasitoid wasps (Chauzat et al 1995 Hanks et al 1995 Rivera et al 2001

Protasova et al 2007) and ladybird beetles (Obrycki and Kring 1998 Baker et al 2003)

Pests which have been successfully controlled include psyllid bugs (Chauzat et al

1995 Dahlsten et al 1998) chrysomelid beetles (Elliot and Greener 1994) the eucalypt

weevil (Gonipterus scuttelatus) (Took 1955 Rivera et al 2001) longicorn beetles

(Hanks et al 1995) leaf blister sawfly (Faulds 1991) and gall wasps (Mendel et al

2007 Protasova et al 2007) One of the main difficulties in achieving successful

biological control is establishing stable populations of the control agent species

(Cameron et al 1993 Rosenheim et al 1999) One solution to this problem is to

periodically release large numbers of predatory species into plantations thereby

inundating pest populations (Baker et al 2003) Although most biological control agents

are specialised predators (Faulds 1991 Elliott and Greener 1994 Eldridge et al 1995

Grundy and Maelzer 2000) some studies show that these are more efficient at pest

control because they alternate between prey items and maintain low but more stable

pests populations (Rosenheim et al 1999) Lacewings ladybird beetles and assassin

bugs may have the potential to be efficient biological control agents in eucalypt

plantations in southern Queensland Assassin bugs in particular appeared to be efficient

predators of chrysomelid larvae A better understanding of the life cycle host

preference and feeding capacity of Pristhesancus sp would be required if it were to be

used as a biological control agent

91

3 An Inventory of Fungal Species Associated with Eucalypt Plantations in Southern Queensland

Introduction

A large number of pathogens have been described from eucalypts in Australia (Dick

1982 Marks et al 1982 Park and Keane 1982a 1982b Lunquist and Purnell 1987

Carnegie et al 1994 Sankaran et al 1995 Carnegie 2000 Keane et al 2000 Park et

al 2000 Carnegie 2002 Maxwell et al 2003 Andjic et al 2007) Although pathogenic

fungi may cause severe damage in eucalypt plantations (Dungey et al 1987 Lundquist

and Purnell 1987 Carnegie et al 1994 Crous and Wingfield 1996) they are also an

integral component of forest ecosystems (Shearer 1994 Sankaran et al 1995 Hansen

1999 Burgess and Wingfield 2002) The majority of pathogens occurring in Australian

plantations are likely to have originated in native forests (Park et al 2000 Strauss 2001

Burgess et al 2006) Pathogens can be accidentally introduced into eucalypt

plantations in association with contaminated germplasm such as seeds seedlings or

soil (Straus 2001) Some of the more common genera of pathogens found in eucalypt

plantations include Quambalaria Teratosphaeria Coniella Harknessia

Cylindrocladium Holocryphia and Neofusicoccum

Quambalaria pitereka is a pathogen which may infect foliage and stems of Corymbia

Blakella Angophora (Walker and Bertus 1971 Bertus and Walker 1974) and

Eucalyptus species (Pegg et al 2008) When affecting foliage the symptoms of Q

pitereka are commonly called Quambalaria shoot blight (Pegg et al 2005 Carnegie

2007b) The pathogen causes necrotic spotting and distortion of young expanding

foliage White spore masses are associated with necrotic lesions which rupture through

the leaf cuticles (Pegg et al 2005) Severe infections may cause shoot dieback stunted

92

growth and death in severe cases (Old 1990 Pegg et al 2005) Although Q pitereka

has been known from nurseries since the late 60s (Walker and Bertus 1971) it has only

recently been found to have an extended host range Susceptible species in tropical

eastern Australia include E grandis and E dunnii (Simpson 2000 Self et al 2002

Pegg et al 2005)

The genus Teratosphaeria contains a large number of ascomycete species which vary

greatly in their pathogenicity on eucalypts (Crous 1997 Corlett 2005 Hunter 2011)

Most species cause necrosis of foliage which ranges in severity from small circular

spots to large irregular leaf blights (Crous 1998) T cryptica is one of the more

pathogenic species and is capable of causing severe damage in plantations (Cheah

1977 Park 1982a 1982b Fry 1983 Crous 1998 Carnegie and Ades 2003 Carnegie

and Keane 2002 Jackson et al 2005) Mycosphaerella species such as M lateralis are

less pathogenic and can be found in association with more pathogenic species

(Jackson et al 2004) E globulus is particularly susceptible to Mycosphaerella species

(Park and Keane 1982a amp 1982 b Carnegie et al 1994 Carnegie et al 1997 Park et

al 2000 Milgate et al 2001 Carnegie amp Ades 2002 Maxwell et al 2003 Mohammed et

al 2003 Milgate et al 2005) E globulus was abandoned as a plantation species in

South Africa because of its susceptibility to Mycosphaerella (now Teratosphaeria)

(Purnell and Lunquist 1986) T cryptica and T nubilosa cause severe damage to E

globulus and E nitens in Australia and South Africa (Crous et al 1989b Carnegie et al

1994 Crous and Wingfield 1996 Dungey et al 1987) In New Zealand T cryptica is

reported to have caused an epidemic which affected over 1000 ha of E delegatensis

(Cheah 1977 Fry 1983)

The genus Teratosphaeria also contains species which were once placed in the genera

Mycosphaerella Phaeophleospora Kirramyces and Colletogloeopsis (Crous et al

93

1989 Crous 1997 Cortinas et al 2006 Andjic et al 2007 Hunter et al 2011) The

three most important Teratosphaeria species in eucalypt plantations are T zuluensis T

destructans and T eucalypti (Wingfield et al 1997 Park et al 2000) T zuluensis

causes stem cankers on eucalypts in sub-tropical climates in many countries around the

world (Winfield et al 1997 Old et al 2003 Cortinas et al 2006 Grezahgne et al

2004 Cortinas et al 2006) T destructans is an aggressive pathogen causing distortion

and blight of foliage buds and shoots in South East Asia (Wingfield et al 1996 Old et

al 2003 Burgess et al 2006) T eucalypti and Readeriella epicoccoides are foliar

pathogens which are endemic to Australia Outbreaks of these species mainly occur in

sub-tropical regions For example in northern New South Wales T eucalypti was found

causing severe damage to E nitens plantations R epicoccoides was found causing

severe damage to E grandis and E grandis x E camaldulensis plantations in northern

New South Wales (Carnegie 2007b) and central and southern Queensland (Pegg et al

2003)

Coniella fragariae is a foliar pathogen of both eucalypt plantations and nurseries The

species has a wide host range and is usually found during the wetter months The

fungus causes necrotic blighting of foliage and is distinct in that its fruiting bodies form

concentric rings within necrotic lesions which are easily recognised (Carnegie 2002)

Interestingly lesions are often associated with insect damage such as that caused by

chrysomelid larvae (Ferreira and Milani 2002)

Cylindrocladium quinqueseptatum is a serious pest of plantations particularly in tropical

regions This species proliferates in wet conditions and is often observed after heavy

rain (Carnegie 2002) Symptoms range from distorted foliage with dark rapidly

expanding lesions to cankers on young stems Severe foliar damage can lead to

premature leaf shedding (Carnegie 2002 Jayasinghe et al 2009)

94

The genus Harknessia contains pathogens which cause shoot diseases of various plant

species More than thirteen species have been found associated with eucalypts from

various parts of the world (Sankaran et al 1995) Only five species have been recorded

in Australia H eucalypti has been found in Western Australia (Sutton 1971 Gibson

1975) the ACT (Yuan 1989) and Tasmania (Yuan and Mohammed 1997b) H

fumaginea has been found in Queensland (Sutton 1975) H uromycoides has been

found in Western Australia (Sutton 1971) H victoriae has been found in Victoria (Sutton

and Pascoe 1989) and H weresubiae has been found in South Australia (Nag Raj

1993) Most of these species were found associated with leaf spots and are not

considered to be aggressive pathogens H eucalypti has been found associated with

stem cankers of eucalypts in eastern Australia (Yuan and Mohammed 1997a)

Eucalypt plantations are also susceptible to a range of canker pathogens Holocryphia

eucalypti is a canker pathogen that has been found causing various levels of damage to

at least 20 species of eucalypts in a range of localities in Australia (Davison 1982

Fraser and Davidson 1985 Walker et al 1985 Old et al 1986 Davison and Coates

1991 White and Kile 1993 Yuan and Mohammed 1997 Wardlaw 1999 Gryzenhout et

al 2006) The species is particularly widespread in the eastern states of Australia where

it is common (Walker et al 1985 Old et al 1986 Yuan and Mohammed 1997 Wardlaw

1999 Carnegie 2007a 2007b) Symptoms vary and may include cracking of the bark

swelling of the stem kino exudation and dieback of coppice shoots branches and

stems (Walker et al 1985 Old et al 1986)

The anamorphs of Botryosphaeria species such as Neofusicoccum ribis may cause a

range of symptoms on eucalypts including dieback stem bleeding necrosis coppice

failure and cankers (Davison and Tay 1983 Smith and Kemp 1994 Old and Davison

2000 Burgess and Wingfield 2002) The species is also an endophyte of healthy hosts

95

but may become pathogenic and cause disease in stressed hosts (latent pathogenicity)

(Old et al 1990 Fisher et al 1993 Zhonghua et al 2001 Burgess et al 2004 Slippers

et al 2004)

Chapter Aim

During disease surveys in plantations in southern Queensland a large diversity of

pathogens were identified The aim of this chapter is to present ecological profiles and

describe the impacts of the more common pathogens identified Taxonomic descriptions

of previously undescribed species are also presented


Site Selection

Twenty eucalypt plantations in southern Queensland were sampled for pathogens

between December 2003 and November 2006 The plantations extended from 60 km

south of Brisbane to 60 km north of Bundaberg The majority of these plantations were

planted with E dunnii which ranged from 1-6 years old Other species which were

sampled to a lesser extent included E grandis E urophylla E tereticormis and E

globulus These plantations also ranged from 1-6 years of age

Sampling Regime

Sampling occurred at three month intervals and lasted 2-3 weeks During each field trip

several plantations in the southern Queensland region were repeatedly sampled while

conducting other experiments (Chapters 4 5 amp 6) Some plantations were sampled

intensely whilst others were sampled opportunistically

Sampling Method

Each plantation was originally sampled over an eight to ten hour period during drive-

96

through surveys (Speight and Wylie 2001) Plantations which had repeated visits were

usually sampled for a further one to two hours on separate field trips thereafter

Sampling was largely opportunistic and involved driving by 4WD vehicle along access

tracks within plantations to several localities and then travelling by foot while searching

for disease symptoms amongst trees Topographical maps were consulted to target

different areas such as flats slopes and hill crests Different soil types were also

targeted (identified in the field) Maximising the sampling effort allowed a potentially

greater number of species to be collected in a short period of time

Two types of diseased material were collected diseased foliage and diseased stems

(cankers) Diseased foliage was removed by hand and placed within paper envelopes

which were then refrigerated below 5 degC Diseased stem material was cut into chips (5 x

5 cm) using a large alcohol sterilised machete and then placed in paper envelopes

which were refrigerated below 5 degC All material was examined microscopically within

two weeks of collection

Fungal Isolation

Isolates were obtained by collecting conidia exuding from single pycnidia using the tip of

a sterile needle These were transferred onto 2 Malt Extract Agar (MEA 20 gL Biolab

malt extract 15 gL Biolab agar) containing streptomycin 150 gml (Sigma-Aldrich

Australia) in a single spot and allowed to hydrate for 5 min Under a dissecting

microscope spores were then streaked using a sterile needle and single spores were

immediately transferred to MEA plates Cultures were grown in the dark at 28C for two

weeks and then transferred to fresh MEA plates All cultures were maintained on 2

MEA in tubes at 20 ordmC

The ascospores of ascomycete species were collected by taping a 1 cm square section

97

of each foliar lesion containing ascocarps to the lid of a Petri-dish containing malt

extract (20 g l-1) agar (MEA) The Petri-dish was placed upside down on a bench and

left overnight to allow the spores to be forcibly ejected onto the media above The

germination pattern of the ejected spores was examined and photographed after 12 hrs

Single germinating spores were then removed from the media using a sterile needle

and placed on fresh media which were maintained in the dark at 20ordm C

Wood chips from diseased stems were cut into smaller pieces under sterile conditions

and then surface sterilised with alcohol and flamed for 2-3 seconds (Old et al 1986)

The pieces were then placed onto Petri-dishes containing (20 g l-1) agar (PDA)

containing streptomycin 150 gml (Sigma-Aldrich Australia) and incubated at 28C for

3-4 days Small pieces of mycelia were then removed from the growing margin of fungal

growths and placed onto fresh Petri-dishes (PDA) which were then maintained in the

dark at 20C

Fungal cultures of any species which were reluctant to produce spores in culture were

placed under mixed light (fluorescent and UV) to encourage sporulation

All isolates are currently maintained in the culture collection at Murdoch University

(MUCC) Reference strains have been deposited in the collection of the Central bureau

voor Schimmel cultures (CBS) Utrecht Herbarium specimens of new collections have

also been lodged in the herbarium of the Murdoch University (MURU) Descriptions

were deposited in MycoBank

Morphological Identification

Disease symptoms of foliage including necrotic lesions and fruiting bodies were

photographed using a Canonreg digital camera (macro setting) The fruiting structures

associated with foliage and those produced in culture were examined at high

98

magnification using a compound microscope (x1000 oil immersion) (Olympus BH2 light

microscopereg) Larger fruiting structures were cut into thin sections by hand using a

small piece of razor blade inserted into a needle holder The sections were then

mounted in both lacto-glycerol and aniline blue solution Smaller fruiting structures were

examined as squash mounts Structures were photographed using a digital camera

(Olympus digital copy) which was mounted on the eyepiece of the compound microscope

Molecular Identification

The isolates were grown on 2 MEA at 20C for 4 weeks and the mycelium was

harvested and placed in a 15 ml sterile Eppendorf reg tube Harvested mycelium was

frozen in liquid nitrogen ground to a fine powder and genomic DNA was extracted A

part of the internal transcribed spacer (ITS) region of the ribosomal DNA operon was

amplified using the primers ITS-1F (5rsquo CTT GGT CAT TTA GAG GAA GTA A) Gardes

and Bruns (1993) and ITS-4 (5rsquo TCC TCC GCT TAT TGA TAT GC 3rsquo) (White et al

1990)

Fungal species were considered to be new if their sequenced amplicons did not match

other species which were lodged with Genbank The morphological characters of the

species were compared with related species described in the literature (especially

those for which sequence data were not available)

Morphological Descriptions

For each undescribed species 5 mm plugs of mycelia were cut from actively growing

cultures and placed at the centres of Petri-dishes (55 mm) containing one of three

different nutrient media The media used were 2 malt extract agar (MEA) oatmeal

agar (OMA 20 g of oats boiled in 1 litre of water 15 g of agar (DNA grade) added and

then autoclaved for 20 min at 120 degC) and eucalypt leaf agar (ELA juvenile E globulus

99

foliage was wet autoclaved for 20 min at 120 degC and then two leaves were placed on

the surface of sterile tap water agar in each Petri-dish) Three replicates of each isolate

were grown on each media type at 28 degC in the dark After 30 days cultures were

assessed for growth and photographed Cultures were measured by taking two

measurements of the colony diameter perpendicular to each other using a 10 mm ruler

Each isolate was assessed for conidial size shape pigmentation and number of septa

Wherever possible 30 measurements (x 1000 magnification) of all taxonomically

relevant structures were recorded for each species and the extremes were presented in

parentheses Munsell soil colour charts were used to describe isolate colouration

(Munsel 1905) Measurements of conidial size were obtained using a graticule eyepiece

in conjunction with a compound microscope using oil immersion (x1000) Structures

were photographed using a digital camera (Olympus) which was mounted to the

microscope Line drawings of conidia and conidiogenous cells were drawn in pencil

using a mounted drawing tube apparatus The drawings were then scanned and

modified using Adobe Photoshop v8 copy program

Phylogenetics

In order to compare Teratosphaeria isolates generated from this study with other closely

related species additional ITS sequences were obtained from GenBank Sequence data

were assembled using Sequence Navigator version 101 (Perkin Elmer) and aligned in

Clustal X (Thompson et al 1997) Manual adjustments were made visually by inserting

gaps where necessary All sequences derived in this study were deposited in GenBank

Parsimony analysis with heuristic search was performed using PAUP (Phylogenetic

Analysis Using Parsimony) (Swofford 2001) with random stepwise addition in 100

replicates with the tree bisection-reconnection branch-swapping option and the

100

steepest-descent option off All ambiguous and parsimony-uninformative characters

were excluded gaps were treated as a fifth character MaxTrees were unlimited

branches of zero length were collapsed and all multiple equally parsimonious trees

saved Estimated levels of homoplasy and phylogenetic signal tree length (TL)

consistency index (CI) and retention index (RI) were determined (Hillis and

Huelsenbeck 1992) Characters were unweighted and unordered branch and branch

node support was determined using 1000 bootstrap replicates with equal probability

(Felsenstein 1985) ITS trees were rooted to Readeriella spp and combined trees

were rooted to Mycosphaerella pini

Bayesian analysis was conducted on the same datasets as the one used in the distance

analysis First MrModeltest v 35 (Nylander 2004) was used to determine the best

nucleotide substitution model Phylogenetic analyses were performed with MrBayes v

31 (Ronquist and Heuelsenbeck 2003) applying a general time reversible (GTR)

substitution model with gamma (G) and proportion of invariable site (I) parameters to

accommodate variable rates across sites Two independent runs of Markov Chain

Monte Carlo (MCMC) using 4 chains were run over 1 000 000 generations Trees were

saved for each 1 000 generations resulting in 1 001 trees Burn-in was set at 100 001

generations (101 trees) well after the likelihood values converged to the stationery

leaving 900 trees from which the consensus trees and posterior probabilities were

calculated The new sequences were deposited in GenBank and the alignments and

phylogenetic trees in TreeBASE (wwwtreebaseorg)

101

Results

Described Fungal Species

A number of pathogens were identified during the survey (Table 31) More than one

species was often isolated from diseased material including saprophytes opportunistic

pathogens and primary pathogens Assemblages of fungi were often isolated from

cankers associated with diseased stems (Figure 3-1)

Species profiles are presented for major pathogens including Readeriella epicoccoides

(Figure 3-2) Mycosphaerella heimii (Figure 3-3) Mycosphaerella marksii (Figure 3-5)

Mycosphaerella lateralis (Figure 3-4) Teratosphaeria nubilosa (Figure 3-6) Readeriella

eucalypti (Figure 3-7) and new Teratosphaeria species (Figure 3-10 Figure 3-11 and

Figure 3-12)

Figure 3-1 Stem canker of a 2-year-old E dunnii host from which Holocryphia eucalypti and Neofusicoccum ribis were both associated A basal canker showing swelling at the base of the stem and cracking of the bark (arrow) B basal canker with bark removed to show necrosis of the vascular cambium (arrow)

102

Species Host Ecology Incidence and Threat

Saprophytic fungi

Alternaria sp E dunnii E grandis E globulus

All saprophytic fungi were found associated with dead plant tissues including foliage and stems Some species were also associated with disease symptoms caused by primary pathogens

Ubiquitous and usually associated with damaged or stressed trees Some species appeared to cause secondary necrosis to foliage already infected with primary pathogens (Cladosporium sp Pestalotiopsis sp being the most common) Low threat

Aspergillus sp

Cladosporium sp

Epicoccum sp

Fusarium sp

Mucor sp

Penicillium sp

Pestalotiopsis sp

Phanerocaeta sordida

Phoma glomerata

Phomopsis diaporthe

Nigrasporum sp

Trichoderma sp

Opportunistic Pathogens

Neofusicoccum ribis E dunnii E grandis

A latent pathogen found associated with stems and foliage in the absence of disease symptoms Often associated with other species such as Cytospora eucalypticola and Holocryphia eucalypti May infect both foliage and stem tissues and may cause stem cankers on stressed hosts Symptoms included dark streaking of the vascular cambium and darkening of the bark surface

Frequently found associated with stem cankers in 1-2-year-old plantations A greater incidence of damage was observed in E dunnii plantations than E grandis plantations although this may have been the result of biased sampling due to greater numbers of E dunnii plantations in the region Low threat

Coniella fragariae E dunnii E grandis

An opportunistic pathogen associated with foliar lesions often in conjunction with insect herbivore damage

Frequently found associated with foliar chrysomelid damage in plantations aged 1-2 years Moderate threat

Cytospora eucalypticola E dunnii E grandis

An opportunistic pathogen often found associated with stem cankers along with other species including Holocryphia eucalypti and Neofusicoccum ribis

Frequently found associated with stem cankers in 1-2-year-old plantations Considered weakly pathogenic although may cause damage in stressed trees Low threat

Table 31 Pathogens found associated with E dunnii E grandis and E globulus plantations in southern Queensland

103



Readeriella eucalypti E dunnii

An opportunistic pathogen often found in association with foliar necrotic lesions caused by primary pathogens such as Teratosphaeria species

Rarely encountered Low threat

Primary Pathogens

Aulographina eucalypti E dunnii E grandis

A primary pathogen associated with foliar necrotic lesions (irregular with distinct elongate fruiting bodies)

Rarely encountered but appeared to be more common on mature foliage of 2-3-year-old plantations Low threat

Cryptosporiopsis sp E dunnii A primary pathogen associated with foliar necrotic lesions (circular distinctly dark and brown)

Only encountered once on a single tree exhibiting negligible damage Low threat

Dichomera sp (Neofusicoccum sp)

E dunnii A pathogen associated with foliar necrotic lesions of foliage also in association with Chalcidoid wasp damage


Holocryphia eucalypti E dunnii E grandis

A primary pathogen often found associated with stem cankers along with other species including C eucalypticola and N ribis

Frequently found associated with stem cankers in 1-2-year-old plantations Capable of causing death to healthy trees High threat

Readeriella epicoccoides E dunnii E grandis

A primary pathogen associated with large necrotic lesions of foliage (usually oozing spore masses are associated with the leaf underside of mature leaves occurring in the lower canopy)

Commonly encountered in plantations Appeared to be more common on E grandis than E dunnii hosts

May eventually cause greater levels of damage if rainfall leads to high humidity allowing greater sporulation spread and infection Moderate threat

104


Primary Pathogens

Mycosphaerella heimii E dunnii A primary pathogen associated with irregular necrotic lesions of foliage (blights and spots)

Abundant in plantations aged 1-2 years Commonly encountered in plantations near Bundaberg but never encountered in plantations around the Brisbane area May eventually cause greater levels of damage if rainfall leads to high humidity allowing greater sporulation spread and infection High threat

Mycosphaerella lateralis E globulus Primary pathogen associated with circular necrotic lesions of foliage (blights and spots)

Rarely encountered and only found affecting E globulus hosts May eventually cause greater levels of damage if rainfall leads to high humidity allowing greater sporulation spread and infection Moderate threat

Mycosphaerella marksii E grandis Primary pathogen associated with circular necrotic lesions of foliage (blights and spots)

Rarely encountered and only found affecting E globulus hosts


Teratosphaeria nubilosa E globulus Primary pathogen associated with circular necrotic lesions of foliage (blights and spots)

Rarely encountered and only found affecting E globulus hosts May eventually cause greater levels of damage if rainfall leads to high humidity and allowing greater sporulation spread and infection Moderate threat

105

Readeriella epicoccoides

Hosts E dunnii and E grandis

Field Symptoms

Readeriella epicoccoides is associated with necrotic foliar lesions which range in

severity from small spots to large blights The underside of the leaf is often covered in

oozing black spore masses while the upper side often produces fewer spore masses

(Figure 3-2A amp B)

Growth on Agar

The growth medium used was (20 g l-1) agar (MEA) containing streptomycin 150 gml

(Sigma-Aldrich Australia) The cultures are generally slow growing dark brown to black

(10YR 21) in colour raised and dense with white aerial hyphae The growing margins

of the culture are usually pale grey before melanising with maturity (Figure 3-2D)

Morphological Characters

The fruiting structures are pycnidial and immersed with a circular ostiole from which

conidia are produced The conidia are slender curved to straight slightly melanised

multiseptate with truncate ends and tapering to a point at the distal end (Figure 3-2E)

Ecology and threat

R epicoccoides has been observed causing severe damage in northern New South

Wales (Carnegie 2007) and moderate levels of damage have been observed in

southern Queensland on a range of species (Pegg et al 2003) During this study R

epicoccoides was only ever observed at low levels on stressed hosts The species

appeared to be more abundant on both E grandis and E grandis x camaldulensis than

E dunnii R epicoccoides is the anamorph of Teratosphaeria suttoniae which was

106

never observed during the study

Given that the drought in southern Queensland has largely abated and R epicoccoides

is known to proliferate in humid conditions (Walker 1962 Chipompha 1987 Walker et

al 1992 Ferreira and Milani 2002) there is potential for an increase in disease The

species is therefore considered to be a moderate threat to the southern Queensland

plantation industry

107

Figure 3-2 Readeriella epicoccoides on E grandis foliage A adaxial leaf surface B abaxial leaf surface (arrows point to oozing spore masses) C cross section of pycnidium associated with leaf containing mature conidia D upper surface of culture on 2 MEA after 21 daysrsquo growth E conidia (arrows) and hyphal fragments from leaf

108

Mycosphaerella heimii

Host E dunnii

Field Symptoms

Mycosphaerella heimii is associated with foliar necrotic lesions which are irregular in

shape with distinct brown raised margins (Figure 3-3A B amp C) The necrosis of the leaf

extends through the leaf lamina Brown immersed fruiting structures occur on both sides

of the lesion but are generally greater in density on the lower surface

Growth on Agar


(Sigma-Aldrich Australia) The cultures were slow growing and formed circular colonies

with concentric rings radiating from the centre (Figure 3-3D) On the surface the rings

range in colour from pale to dark browngreen (10YR 24) with pale aerial hyphae The

outer margin is pale before melanising with maturity The underside of the culture is

darker than the upper surface and brown to black (10YR 11)


The ascocarps are flask-shaped and immersed within the leaf tissue The asci have a

distinct foot attachment at the base (Figure 3-3F) The germination pattern of the

ascospore after 24 hrs consists of multibranching germtubes from either end of the

spore (Figure 3-3G)

Ecology and Threat

M heimii was the most commonly identified Mycosphaerella species in southern

Queensland The severity of damage varied greatly between trees and was most

abundant during early summer The most severe infection was observed within a two-

109

year-old E dunnii plantation in which a small group of trees had damage to

approximately 70 of their canopies

Given that M heimii was observed to be capable of causing severe damage the

species is considered to pose a high threat to the southern Queensland plantation

industry Trees exhibiting symptoms should be monitored closely and in severe cases

should be removed to reduce further spread of the disease Chemical spraying is only

likely to be effective if the infection to be treated is highly localised

110

Figure 3-3 Teratosphaeria heimii on E dunnii foliage A adaxial leaf surface (arrows point to lesions) B abaxial leaf surface (arrows point to lesions) C lesions with raised brown margins D upper surface of culture on 2 MEA after 21 daysrsquo growth E lower surface of culture on 2 Malt Extract Agar after 21 daysrsquo growth F asci with distinct foot attachment (arrow A) and containing ascospores (arrow B) G germination pattern of ascospore after 24 hrs on 2 Malt Extract Agar arrow points to germinating hyphae

A

B

111

Mycosphaerella lateralis

Host E globulus

Field Symptoms

Mycosphaerella lateralis is associated with foliar necrotic lesions which are irregular in

shape with distinct red brown margins (Figure 3-4A B amp C) Necrosis of the leaf extends

through the leaf lamina Brown immersed fruiting structures occur on both sides of the

lesion but are generally denser on the lower surface

Growth on Agar


(Sigma-Aldrich Australia) The cultures are slow growing and form a circular colony with

pale aerial hyphae and a distinct yellow margin (10YR 62) in the agar (Figure 3-4D)

The underside of the culture is darker than the upper surface and green brown (10YR

24) the yellow growth margin is also visible (Figure 3-4E)


The ascocarps are flask-shaped and immersed within the leaf tissue The germination

pattern of the ascospore after 24 hrs consists of lateral germ tubes emerging from either

end of the spore (Figure 3-4F)

Ecology and Threat

M lateralis was found causing low levels of damage to a four-year-old stand of E

globulus The literature suggests that the species may be a parasite of other

Teratosphaeria species (Jackson et al 2004) T nubilosa (a known primary pathogen)

was also isolated from lesions in association with M lateralis which supports this

hypothesis Given that only low levels of the disease were observed (no damage was

112

observed within E dunnii plantations) M lateralis is considered to be a low threat to the

plantation industry in southern Queensland

Figure 3-4 M lateralis on E globulus foliage A adaxial leaf surface (arrows point to lesions) B abaxial leaf surface (arrows point to lesions) C circular lesion with red-brown margin D upper surface of culture on 2 MEA after 21 daysrsquo growth E lower surface of culture on 2 Malt Extract Agar after 21 daysrsquo growth F germination pattern of ascospore after 24 hrs on 2 Malt Extract Agar

113

Mycosphaerella marksii

Host E grandis

Field Symptoms

Mycosphaerella marksii is associated with foliar necrotic lesions which are pale brown

irregular in shape with distinct red brown margins (Figure 3-5A B amp C) Necrosis of the

leaf extends through the leaf lamina Brown immersed fruiting structures occur on both

sides of the lesion but are generally greater in density on the lower surface

Growth on Agar


(Sigma-Aldrich Australia) The cultures are slow growing and form circular colonies with

pale aerial hyphae on the surface (10YR 62) (Figure 3-5D) The underside of the

culture is darker than the upper surface and brown to black (10YR 34) (Figure 3-5E)



pattern of the ascospore after 24 hrs consists of a single germ tube which emerges

perpendicular to the ascospore and is very long (Figure 3-5F)

Ecology and Threat

M marksii was found causing low levels of damage to a four-year-old stand of E

grandis The incidence of the disease appeared to be relatively constant and did not

appear to vary greatly between winter and summer

Given that only low levels of the disease were observed (no damage was observed

within E dunnii plantations) M marksii is considered to be a low threat to the plantation

114


Figure 3-5 Mycosphaerella marksii on E dunnii foliage A adaxial leaf surface (arrows point to lesions) B abaxial leaf surface (arrows point to lesions) C lesions with red-brown margins D upper surface of culture on 2 MEA after 21 daysrsquo growth E lower surface of culture on 2 Malt Extract Agar after 21 daysrsquo growth F germination pattern of ascospores after 24 hrs on 2 Malt Extract Agar

115

Teratosphaeria nubilosa

Host E globulus

Field Symptoms

Teratosphaeria nubilosa is associated with foliar necrotic lesions which are light brown

circular in shape with distinct red brown margins (Figure 3-6A B amp C) The necrosis of

the leaf extends through the leaf lamina Brown immersed fruiting structures occur on

both sides of the lesion but are generally greater in density on the lower surface

Growth on Agar


(Sigma-Aldrich Australia) Cultures are slow growing and form irregular shaped

colonies with pale aerial hyphae on the surface (10YR 62) (Figure 3-6D) The outer

margin of the colony is pale green (10YR 34) The underside of the culture is darker

than the upper surface and brown to black (Figure 3-6E)


The ascocarps are flask shaped and immersed within the leaf tissue The germination

pattern of the ascospore after 24 hrs consists of two short germ tubes extending parallel

from either end of the ascospore (Figure 3-6F)

Ecology and Threat

T nubilosa was one of the most abundant Teratosphaeria species in E globulus

plantations in southern Queensland The species was frequently associated with foliar

damage in a four-year-old stands of E globulus but was never found associated with E

dunnii T nubilosa was isolated from the same lesions as M lateralis which may

indicate that they have a hyper-parasitic relationship Because M nubilosa was not

116

found associated with E dunnii the species is considered to pose a low threat to the


Figure 3-6 Teratosphaeria nubilosa on E globulus foliage A adaxial leaf surface (arrows point to lesions) B abaxial leaf surface (arrows point to lesions) C circular lesion with dark brown margin D upper surface of culture on 2 MEA after 21 daysrsquo growth E lower surface of culture on 2 Malt Extract Agar after 21 daysrsquo growth F germination pattern of ascospore after 24 hrs on 2 Malt Extract Agar

117

Readeriella eucalypti

Host E dunnii

Field Symptoms

Readeriella eucalypti is associated with foliar necrotic lesions which are irregular pale

brown with raised dark brown margins (vein limited) (Figure 3-7A B F amp G) Fruiting

structures are ascervular (Figure 3-7I) which vary in size and have distinctly narrow

ostioles

Growth on Agar


(Sigma-Aldrich Australia) A fast growing pale cream-white culture with white aerial

hyphae on the surface (10YR 81) (Figure 3-7C E amp H) The lower surface is dark

brown (10YR 24) (Figure 3-7D)


Conidiomata Pycnidial globular ostiolate superficial 3-7 layers of textura angularis

with conidiogenous cells attached to the inner wall (Figure 3-7I) Conidiogenous cells

Hyaline ellipsoidal to subovate when produced becoming melanised and globular with

flat edges and a marginal frill (Figure 3-7J) Conidia holoblastic melanised globular

thick walled with an acute tip at base (Figure 3-7J)

Ecology and Threat

Given that R eucalypti was only isolated on a single occasion from one E dunnii host

the species is considered to pose a low threat to the plantation industry in southern

Queensland

118

Figure 3-7 Readeriella eucalypti on E dunnii foliage A adaxial leaf surface (arrows point to lesions) B abaxial leaf surface (arrows point to lesions) C upper surface of culture on 2 MEA after 21 daysrsquo growth D lower surface of culture on 2 Malt Extract Agar after 21 daysrsquo growth E upper surface of culture on Oatmeal agar after 21 daysrsquo growth F lesion with a brown raised margin G Pycnidia associated with lesion surface (arrows) H upper surface of culture on Eucalypt Leaf Agar after 21 daysrsquo growth I mature pycnidium oozing conidia J conidiogenesis of immature hyaline conidia and mature melanised conidia as on leaf

119

New Fungal Species

Three new species of Teratosphaeria were collected A BLASTn search was first

conducted on GenBank to compare the ITS sequences of the new Teratosphaeria spp

with those lodged in GenBank Other Teratospheria species known from eucalypts were

also used in a phylogenetic analysis (Figure 3-) TreeBASE SN4443) The aligned ITS

dataset contained 553 characters of which 203 were parsimony informative and

contained significant phylogenetic signal (Plt001 g1=-139) Parsimony analysis

resulted in 34 most parsimonious trees of 679 steps (CI=051 RI=083) Whilst there is

strong bootstrap and Bayesian support for terminal species clades and for some groups

of species there is little support for higher order clustering T micromaculata sp nov

and T biformis sp nov cluster together separate from other Teratopshaeria species

separated from each other with high bootstrap and Bayesian support T aurantia sp

nov also resides in a strongly supported terminal clade clustering with T syncarpiae

and T fibrillossa (Figure 3-8)

Taxonomic descriptions are presented for the three new species of Teratosphaeria

(Table 32)

120

Teratosphaeria nubilosa CMW11560 DQ658232

Teratosphaeria nubilosa CBS114708 AF449099

Teratosphaeria eucalypti CMW17917 DQ632711

Teratosphaeria eucalypti CBS113992 DQ240001

Teratosphaeria destructans CMW17918 DQ632666


MUCC467 EU300999

MUCC468 EU301000

MUCC649 DQ240133

MUCC693 EU301002

MUCC694 DQ240169

Teratosphaeria veloci CPC14600 FJ023539

Teratosphaeria cryptica CBS110975 AY309623

Teratosphaeria cryptica MURU115 AY509754

Teratosphaeria suttonii MUCC425 DQ632655

Teratosphaeria corymbiae CBS120495 EF011657


Teratosphaeria toledana CPC10840 AY725581

Teratosphaeria toledana CBS113313 AY725581

Teratosphaeria callophylla MUCC700 FJ641060


Teratosphaeria pseudocryptica CPC11264 DQ303009

Teratosphaeria pseudocryptica CBS118504 DQ303010

Teratosphaeria rubidae MUCC659 FJ532013


Teratosphaeria fimbriata CPC13321 EF394835

Teratosphaeria angophorae CBS120493 EF011653


Teratosphaeria tinara MUCC665 EU300993



Terarosphaeria tinara MUCC665 EU300997

Teratosphaeria multiseptata DAR77440 DQ530223


Teratosphaeria limosa MUCC695 FJ532010


MUCC668 EU301011

MUCC669 EU301014

Teratosphaeria syncarpiae DAR77433 DQ530219

Teratosphaeria syncarpiae NSWF005320 DQ530220

Teratosphaeria fibrillosa CBS121707 EU707862

Teratosphaeria fibrillosa CPC13969 EU707863

Teratosphaeria dimorpha CBS120085 DQ923529

Teratosphaeria pluritubularis CBS118508 DQ303007

Teratosphaeria ovata CPC14632 FJ023538

Teratosphaeria brunneotingens CPC13303 EF394853

Teratosphaeria molleriana CBS117924 DQ239968

Teratosphaeria molleriana CBS111164 AF309620

Teratosphaeria molleriana CBS110499 AY150675

Teratosphaeria stellenboschiana CBS116428 AY725518

Teratosphaeria gauchensis CBS117257 DQ240198


Teratosphaeria foliensis MUCC670 EU301006


Teratosphaeria zuluensis CBS117835 DQ239987


Teratosphaeria considenianae CBS120087 DQ923527

Teratosphaeria blakelyi CBS120089 DQ923526

Teratosphaeria juvenalis CBS110906 AY725513


Teratosphaeria verrucosa CPC18 AY725517

Teratosphaeria verrucosa CBS113621 AY725515

Readeriella novaezelandiae CBS114357 DQ267603

Readeriella novaezelandiae CPC10895 AY725578

Readeriella mirabilis CPC10506 AY725529

Readeriella mirabilis CPC11712 DQ303094

Readeriella readeriellophora CPC10375 AY725577

Readeriella readeriellophora CPC11711 DQ303013

Readeriella eucalypti CPC11735 DQ303093


5 changes

100

67

100

100

76

100

75

96

100

55

100

100

100

99

92

85

87

57

100100

99

100

100

100

100

97

100

100

86

98

66

97

91

97

100

84

84

99

98

88

52

Teratosphaeria aurantia

Teratosphaeria biformis

Teratosphaeria micromaculata

Figure 3-8 Parsimony analysis resulting in 34 most parsimonious trees of 679 steps (CI=051 RI=083) Each of the new Teratosphaeria species are highlighted in grey

121

Species Hosts Ecology and Field Symptoms Incidence and Threat

Teratosphaeria aurantia sp nov E dunnii (4-year-old)

E grandis (4-year-old)

A primary pathogen associated with foliar necrotic lesions

Lesions small to moderate circular pale brown with a dark brown margin usually with a distinct aggregation of black fruiting bodies near the lesion centre (Figure 3- F amp G)

Lesions scattered over the leaf and extending through the leaf lamina (Figure 3- A amp B)

Localised incidence (20-30 trees in small areas) Usually with a large proportion of leaves affected on each host (40 ) Moderate threat

Teratosphaeria biformis sp nov E dunnii (4 ndashyear-old) E globulus (3-year-old)

A primary pathogen associated with foliar necrotic lesions Sometimes found associated with the same lesions as K aurantia sp nov

Lesions small to moderate circular or irregular pale in colour with a raised purple margin

Lesions scattered over the leaf and extending through the leaf lamina


Teratosphaeria micromaculata sp nov

E globulus (3-year-old) A primary pathogen associated with foliar necrotic lesions

Lesions small circular dark brown and raised


Localised incidence (10-20 trees in small areas) Low threat

Table 32 New Teratosphaeria species found associated with E dunnii E grandis and E globulus (December 2003 and November 2006)

122

Teratosphaeria aurantia sp nov

Etymology named after the orange colour of the cultures

Taxonomic Description

Leaf spots epiphyllous and hypophyllous extending through leaf lamina light brown

conspicuously circular 05-5 mm diameter (Figure 3-10 A amp B) with corky brown

margins (Figure 3-10 F) Mycelium immersed in host tissue septate branching

melanised Conidiophores reduced to conidiogenous cells (Figure 3-10 J)

Conidiomata pycnidial sub-epidermal separate globose wall of 4-5 layers of dark

brown textura angularis (Figure 3-10 I) Conidiogenous cells sub-cylindrical sub-

hyaline to medium brown smooth proliferating percurrently and enteroblastically with 1-

4 annulations formed from the inner cells of the pycnidial wall 55 x 40 μm (Figure 3-10

J) Conidia ellipsoidal 0-1 septate subhyaline to medium brown smooth eguttulate

falcate gradually tapering toward apex truncate at base (95ndash)11-14(ndash160) x (25ndash)25-

35(ndash40) (mean = 125 x 30 μm (Figure 3-10 J)

Cultural characteristics Colonies on MEA reaching diam 4 x 5 mm after 1 month at 28

C globular aggregating or separate masses with white to cream (2Y 883) short aerial

hyphae on the surface dark brown (10YR 48) on reverse (Figure 3-10 C amp D) On OMA

colonies reaching 7 x 8 mm diam after 1 month globular aggregating or separate

masses with white to cream (2YR 883) short aerial hyphae on surface dark brown

10YR 33 on reverse (Figure 3-10 E)

Material examined Australia Queensland Rosedale on leaves of E grandis (G

Whyte 2007) holotype MURU440 culture ex-type MUCC668 Additional specimens

Australia Queensland Rosedale on leaves of E dunnii G Whyte 2007 (MURU439)

(culture ex-type MUCC669)

123

Notes Although phylogenetically distinct Teratosphaeria aurantia is morphologically

similar to T pseudocryptica and T rubidae However it can be distinguished from the

latter species by the golden yellow stain of agar (T rubidae produce reddish stains on

agar) and slightly thinner conidia (11-14 x 25-35 μm) than Teratosphaeria

pseudocryptica (12-14 x 4 μm) and T rubidae (125-13 x 55-60 μm) In addition T

aurantia lesions are distinctly circular with raised margins and an aggregation of fruiting

structures in the centre

124

Figure 3-10 Teratosphaeria aurantia sp nov on E dunnii foliage A adaxial leaf surface (arrows point to lesions) B abaxial leaf surface (arrows point to lesions) C upper surface of culture on 2 MEA after 21 daysrsquo growth D lower surface of culture on 2 Malt Extract Agar after 21 daysrsquo growth E upper surface of culture on Oatmeal agar after 21 daysrsquo growth F circular lesion with a brown raised margin G Pycnidia associated with lesion surface (arrows) H upper surface of culture on Eucalypt Leaf Agar after 21 daysrsquo growth I mature pycnidium containing mature conidia J conidiogenesis of conidia and detached conidia as on leaf

125

Teratosphaeria biformis sp nov

Etymology named after its ability to produce conidia both as a coelomycete on the leaf

and as a hyphomycete on agar


Leaf spots epiphyllous and hypophyllous light brown conspicuously circular 05-5 mm

diameter extending through leaf lamina (Figure 3-11 F amp G) Mycelium immersed in

host tissue septate branching melanised Conidiophores absent Conidiomata

pycnidial dark brown amphigenous aggregated globose (Figure 3-11 I)

Conidiogenous cells subcylindrical pale brown to brown smooth proliferating

percurrently Conidia holoblastic melanised ovoid thick walled truncate at base (-60)

7-10(ndash110) x (25ndash) 3-4 (ndash40) (mean = 85 x 35 μm) (Figure 3-11 J amp K)

Cultural characteristics Colonies on MEA reaching diameter 30 x 35 mm after 1 month

at 28 C irregular with smooth margins white to cream 2Y 883 short aerial hyphae on

top reverse dark brown with paler brown 10YR 33 83 margins (Figure 3-11 C amp D)

On OMA colonies reaching 60 x 65 mm diameter irregular with smooth margins white

to cream 2Y 883 mycelia with short aerial hyphae on top not visible on reverse (Figure

3-11 E)

Material examined Australia Queensland Rosedale on leaves of E globulus (G

Whyte 2007) MURU438 culture ex-type MUCC693 Additional specimens Australia

Queensland Rosedale on leaves of E dunnii (G Whyte 2007) (MURU435) (culture

ex-type MUCC649)

Notes T biformis is phylogenetically closest to T micromaculata from which it differs by

slightly longer and wider conidia (7-10 x 3-4 μm) compared with T micromaculata (5-7 x

2-3 μm) T biformis is morphologically closest to T ovata but it can be distinguished by

126

its faster growth in culture on MEA (T biformis=35 mm T ovata=20 mm) and OMA (T

biformis=65 mm T ovata 30 mm) It is also ecologically different to other

Teratosphaeria species in that it is one of few described species known to produce

conidia both as a coelomycete in vivo and as a hyphomycete in vitro

Figure 3-11 Teratosphaeria biformis sp nov on Eucalyptus dunnii foliage A adaxial leaf surface (arrows point to lesions) B abaxial leaf surface (arrows point to lesions) C upper surface of culture on 2 Potato Dextrose Agar after 21 daysrsquo growth D lower surface of culture on 2 Potato Dextrose Agar after 21 daysrsquo growth E upper surface of culture on Oatmeal agar after 21 daysrsquo growth F circular lesion with a purple raised margin G spore masses associated with lesion surface (arrows) H upper surface of culture on Eucalypt Leaf Agar after 21 daysrsquo growth I cross-section of spore mass J conidiogenesis from conidiogenous cells as on leaf K conidiogenesis from hyphae as in culture

127


Etymology named after its association with relatively small lesion spots


Leaf spots epiphyllous and hypophyllous dark brown circular lesion 05-2 mm

diameter with a raised purple margin followed by a light brown margin extending

through leaf lamina (Figure 3-12 A amp B) Mycelium immersed in host tissue septate

branching melanised Conidiophores absent Conidiomata acervular globular

superficial with very little of the epidermis remaining intact conidiogenous cells attached

at base (Figure 3-12 J) Conidiogenous cells globular to dolliform medium brown

smooth proliferating percurrently (40ndash) 48 (ndash56) x (40ndash) 45 (ndash48) (Figure 3-12 K)

Conidia ellipsoidal ovoid thick walled guttulate hyaline when produced but becoming

melanised truncate at base (50ndash) 5-7 (ndash75) x (20ndash) 2-3 (ndash35) (mean = 60 x 25 μm)

(Figure 3-12 K)


C irregular with smooth margins dark olive brown 25Y 33 with darker margins light

olive brown 25Y 54 aerial hyphae (Figure 3-12 C amp D) On OMA colonies reaching 12

x 15 mm light olive brown 25Y 54 mixed with light cream hyphae rough lightly furred

(Figure 3-12 E amp F)

Material examined Australia Queensland Boonah on leaves of E globulus (G Whyte

2007) holotype MURU437 culture ex-type MUCC647 Additional specimens Australia

Queensland Boonah on leaves of E globulus (G Whyte 2007) (culture ex-type

MUCC648)

Notes T micromaculata is phylogenetically closest to T biformis but differs by slightly

smaller conidia (5-7 x 2-3 μm) than T biformis (7-10 x 3-4 μm) Morphologically T

128

micromaculata is somewhat similar in conidial shape and size to T gauchensis (5-6 x

25 μm) However it can be easily distinguished from T gauchensis by its lack of

conidiophores as it produces conidia directly from conidiogenous cells

129

Figure 3-12 Teratosphaeria micromaculata sp nov on E dunnii foliage A adaxial leaf surface (arrows point to lesions) B abaxial leaf surface (arrows point to lesions) C upper surface of culture on 2 Malt Extract Agar after 21 daysrsquo growth D lower surface of culture on 2 Malt Extract Agar after 21 daysrsquo growth E upper surface of culture on 2 Potato Dextrose Agar after 21 daysrsquo growth F lower surface of culture on 2 Potato Dextrose Agar after 21 daysrsquo growth G circular lesion with raised brown and purple margins H spore masses associated with lesion surface I upper surface of culture on Eucalypt Leaf Agar after 21 daysrsquo growth J cross section of spore masses associated with lesion surface K conidiogenesis from conidiogenous cells as on leaf

130

Discussion

Twenty-nine species of fungi were identified during the survey These included thirteen

saprophytic or weakly pathogenic species four opportunistic pathogens and twelve

primary pathogens (including three new species) It is expected that these species

represent a small fraction of the diversity of fungi which are likely to occur in plantations

in southern Queensland This is mainly because sampling coincided with a period of

severe drought (2003-2006) which was likely to have adverse effects on many fungal

species

Saprophytes and Drought

It is likely that the drought may have favoured some fungal species such as those

which exploit stressed and dead hosts Thirteen saprophytic or weakly pathogenic

species were isolated from the necrotic tissues of diseased trees Due to the high

incidence of wilting caused by the dry conditions the greater availability of necrotic

tissue in plantations may have also benefited saprophytic species Some species

previously thought to be saprophytic such as Pestalotiopsis sp were isolated from hosts

exhibiting symptoms typical of a primary pathogen These hosts were severely stressed

and may have had reduced resistance Inoculating healthy hosts under controlled

conditions would help elucidate the pathogenicity of these species

Foliar Pathogens and Drought

Foliar pathogens may be negatively impacted by drought conditions because many

species depend on high humidity for sporulation Rainfall is also important for lsquosplash

dispersalrsquo of fungal spores (Howe 1955 Walklate et al 1989 Daniel and Shen 1991)

Leaf wetness has been shown to increase the rate of infection by foliar pathogens

(Beaumont 1947 Krausse and Massie 1975) During severe drought it was observed

Asci

131

that premature leaf loss occurred on stressed hosts This may lead to reduced inoculum

levels of pathogens within tree canopies (Figure 3-13) Although many pathogens

sporulate on dead leaves foliage on the ground is likely to disseminate fungal spores to

a lesser extent than canopy foliage

It is likely that if conditions had been more typical of the subtropical climate in southern

Queensland some of the more common fungal species may have been found in

plantations For example Teratosphaeria cryptica is one of the most common foliar

pathogens in eucalypt plantations in eastern Australia (Park and Keane 1982 Crous

and Wingfield 1996 Park et al 2000) This species was never collected in plantations

in southern Queensland

Opportunistic Pathogens and Drought

Opportunistic pathogens such as Neofusicoccum Holocryphia and Cytospora species

are often thought to be ubiquitous in plantations (Old et al 1990 Fisher et al 1993

Yuan and Mohammed 1997) These species were found associated with basal cankers

in one and two-year-old plantations The incidence of Holocryphia eucalypti appeared to

decrease as the drought continued This may indicate that although host stress may

Figure 3-13 Accumulated dead foliage on the ground beneath a stressed E dunnii host suffering premature leaf loss B a fallen leaf with associated lesions (arrows) likely to be caused by a foliar pathogen which affected the leaf while it was alive on its host

132

benefit H eucalypti once it infects its host excessively dry climatic conditions may have

adverse affects on the fungal life cycle outside the host (spore survival dispersion

germination and host penetration) This has been suggested by some authors for other

pathogens (Walker and Stahmann 1955 Cook and Papendick 1972)

New Pathogenic Species

Three new species of Teratosphaeria were identified It is difficult to determine if the

new species pose a threat to the plantation industry because the hosts from which the

species were collected were severely moisture stressed Conversely if the climate in

southern Queensland returns to more typical subtropical conditions (higher humidity)

this may cause an increase in the incidence of these species Given that all new

species were locally restricted at the time of their collection it would be interesting to

examine how these may spread within and between plantations during optimal climatic

conditions

T micromaculata sp nov was only found associated with foliage of E globulus and may

not include E dunnii within its host range T aurantia sp nov and T biformis sp nov

were both isolated from more than one host species which may suggest that they have

a greater host range A better understanding of the pathogenicity of these species

would require a pathogenicity experiment under controlled conditions such as in the

glasshouse

Controlling Pathogens

Pathogens are most commonly controlled in plantations by selectively breeding

plantation trees for greater resistance (Arnold et al 1998) Fungicides are rarely used to

reduce outbreaks of pathogens because fungal spores are generally ubiquitous and can

survive in refugia such as leaf litter (Dickman 1992) Chemical control is often effective

133

in the nursery under controlled conditions

Selective breeding plantation trees for greater resistance to pathogens involves

screening large numbers of trees in the nursery and then propagating the most resistant

varieties (Alfenas et al 1983 Denison and Kietzka 1993 Dianese et al 1984

Gryzenhout et al 2003) Given that disease resistance is often controlled by a limited

number of plant genes selective breeding is often limited to developing resistance to

single species of pathogens (Keen 1990)

Maintaining good plantation hygiene can also reduce the spread of pathogens in

plantations and may involve removing dead branches from unhealthy trees or removing

entire trees with disease symptoms Infected trees are a source of inoculum which can

lead to further spread of disease Simple cultural practices have been shown to be

effective for controlling pathogens such as Armillaria spp which require specific

conditions for infection such as extended periods of high soil moisture or host wounding

(Hickman and Perry 1997 2003)

Conclusion

The drought in southern Queensland (2003-2006) had a negative impact on the majority

of the pathogens found in plantations however some saprophytes and opportunistic

pathogens may have benefited from host stress These species were observed in some

cases causing more severe levels of damage An examination of plantations under

more typical climatic conditions is likely to result in the identification of a number of

pathogenic species not previously encountered

134

4 Pests and Pathogens of Eucalypt Plantations in Southern Queensland Effects of Plantation Age Local Climate and Season

Introduction

The eucalypt plantation industry in southern Queensland is in its infancy and the

ecologies of many pests and pathogens are poorly understood Most strategies to

control pests and pathogens in southern Queensland have been adopted from those

used in other Australian plantation centres

In 2003 research was instigated to provide information about pests and pathogens of

eucalypt plantations in southern Queensland to the plantation industry The effects of

plantation age local climate and season were identified as key areas of research to be

addressed Examining these effects would allow a greater understanding of the

conditions suitable for outbreaks of pests and pathogens

The Effects of Plantation Age

The age of plantation trees may influence the abundance of pests and pathogens in

plantations (Carne 1974) This is mainly because as eucalypts mature the physical and

chemical characteristics of their foliage often changes (Lowman 1984 Zanuncio et al

1998) Stone (1991) in a discussion paper listed a number of important pests of

plantations which prefer either young or mature plantations and suggested that

defoliators prefer young plantations with open canopies while borers prefer mature

plantations with a closed canopy

Many eucalypt species have different forms of juvenile and adult foliage (Heteroblasty)

For example the juvenile leaves of many species are larger softer and more glaucous

than adult foliage (Day 1998 Brennan and Weinbaum 2001) Some pests prefer

135

juvenile eucalypt foliage to adult foliage (Macauley and Fox 1980 Larson and Ohmart

1998 Steinbauer et al 1998 Brennan and Weinbaum 2001 Lawrence et al 2003) In

plantations this trend is particularly strong in chrysomelid beetles and many species

prefer juvenile foliage or new growth instead of adult foliage (Tanton and Khan 1978)

Juvenile foliage also contains less phenolic compounds and has greater available

nitrogen and insects often target this foliage for its greater nutritional value (Landsberg

1990a Kavanagh and Lambert 1990 Abbott et al 1993)

Differences in susceptibility to pathogens also occurs between adult and juvenile

eucalypt foliage For example the juvenile foliage of E globulus has been found to be

more susceptible to infections by Teratosphaeria leaf blight than mature foliage

(Carnegie et al 1994 Andjic et al 2007)

E dunnii plantations tend to have canopies consisting entirely of juvenile foliage for the

first 1-2 years after which they begin to produce mature foliage (pers obs) This would

suggest that younger plantations are more likely to have a greater incidence of pests

and diseases than older plantations however other factors such as the rate of

colonisation (either from native forests or neighbouring plantations) may also have an

influence The lsquohoneymoon periodrsquo predicts that newly established plantations have a

lower incidence of pests and pathogens (Burgess and Wingfield 2002)

The Effects of Local Climate

The southern Queensland region is approximately 61 million ha and climate is variable

across this area From the coast to the interior there is a general trend of decreasing

rainfall and increasing temperature From north to south there is a general trend of

decreasing temperature and rainfall (BOM) Other factors such as topography may also

affect local climate (Hammer 2000)

136

The worldsrsquo insect diversity is concentrated in the tropics and subtropics (Stork 1988)

Insects can proliferate in such climates because high temperatures tend to accelerate

egg and larval development This can increase the chances of survival by reducing the

time spent in the development stages which are more susceptible to predation and

parasitism This can also lead to additional generations per year (Anilla 1969

Yamamura and Kiritani 1998 Wermelinger 2004) High humidity can also benefit

insects by reducing fatality from dehydration (Anilla 1969 Wermelinger 2004) Greater

potential for pest outbreak is one of the main reasons plantation growers in Australia

have avoided tropical sites for growing eucalypt plantations (Carnegie et al 2005)

Diversity determined by climate also occurs in fungal communities Fungi are abundant

in the tropics (particularly pathogens and saprobes) (Van der Kamp 1991 Kendrik

1992) High temperatures and extended periods of leaf wetness of the host can allow

greater rates of sporulation dispersion hyphal development and penetration into host

tissues (Beaumont 1947 Krausse and Massie 1975)

In southern Queensland where the climate is increasingly tropical at higher latitudes it

is expected that plantations occurring at higher latitudes would be exposed to a more

subtropical climate Therefore it is expected that a greater diversity of pests and

pathogens would occur in plantations in the northern region

The Effects of Season

The abundance of many pests and pathogens of eucalypt plantations are seasonally

dependent Species with univoltine life cycles may be attuned to seasonal conditions

and may have specific stages of development which coincide with specific seasons

(Mathews 1976) For example the eggs of many moth species will overwinter in a

suspended state (often under bark or leaf litter) before emerging as larvae in spring to

137

feed on plant hosts during the warmer months The larvae pupate early in early summer

and lay eggs before winter (Common 1970) The larval stages of several Lepidoptera

species are pests of plantations (Heather 1975 McQuillan 1985 Nielsen 1986 Farr

2002) Season can also influence tree recovery following insect attack For example

energy reserves of some tree species may be low at the end of summer after a period

of rapid growth (Stone 1991)

The susceptibility of eucalypts to pathogens can also vary between seasons (Shearer et

al 1987 Tippett et al 1987 1989) Most pathogens only sporulate during periods of

high humidity and rain which may facilitate the spread of spores by splash dispersal

(Walklate et al 1989) For this reason prolonged wet conditions can allow the spread of

fungal diseases (Luque et al 2002) Given that high temperatures and rainfall coincide

during the summer months in southern Queensland this would suggest that pathogens

would be most prevalent during such periods However it is important to note that

climate can also influence the health of plantation trees which may be favoured by high

temperatures and rainfall Host vigour has been shown to sometimes ameliorate pest

and pathogen impacts (Benson and Hager 1993 Stone 2001)

Chapter Aim

Although the effects of plantation age local climate and season on pests and pathogens

have been examined in previous studies in other parts of the world no research has

directly examined these effects in E dunnii plantations in southern Queensland Without

such research only the most tentative assumptions about the dynamics of pests and

pathogens in plantations can be made

The aim of this chapter was to monitor the incidence and severity of target pests and

pathogens in variously aged plantations which occur in two different regional climates of

138

southern Queensland (north and south) Monitoring was conducted throughout a 12

month period so that seasonal effects could also be examined

Materials and Methodology

Site Selection

Eight E dunnii plantations were selected for the study These plantations consisted of

trees which were sourced from the same nursery stock (propagated from local seed)

Four of the plantations occurred approximately 50 km south of Brisbane (Southern

plantations) These plantations were aged approximately one two three and four years

at the beginning of the study Each of the southern plantations occurred within a 10 kmsup2

radius (Figure 4-1) A second age series of plantations also one two three and four

years old were selected approximately 60 km north of Bundaberg (northern

plantations) These plantations also occurred within a 10 kmsup2 radius The northern and

southern plantation groups were separated by over 360 km (Figure 4-1)

The northern and southern groups of plantations were selected because they had

similar site characteristics (Table 41 Table 42) All plantations were partially

surrounded by mixed agricultural land and remnants of native vegetation The

topography of the plantations varied although most occurred in gradual undulating

terrain Prior to plantation establishment all sites previously supported pasture for

horsesgrazing cattle Small portions of remnant vegetation were left intact within some

plantations (particularly within drainage lines) All plantations were considered to have a

history of lsquogood healthrsquo with no previous outbreaks of pests or diseases The two groups

were also compatible in that they occurred at similar distances from the coast (gt50 km)

The main difference between the plantations was their respective ages (1-4 years) and

their respective regions (north and south)

139

Table 41 Site characteristics of the southern plantation group Group - Age

Size (ha)

Topography Remnant Vegetation Soil Type Clearing and Fertiliser History

Southern - 1 4355 Gradually sloping with an east west aspect No gullies or deep drainage lines

The entire site was previously used for grazing pasture which was dominated by native grasses and couch Remnant vegetation is unknown but surrounding areas are dominated by E tereticornis and E crebra

Granite based clay loam duplex soil

Progressive clearing since settlement Fertiliser history has been inconsistent

Southern - 2 226 Slightly undulating with gradual slopes and shallow creek lines

Most of the site was previously used for grazing pasture which was dominated by native grasses and couch Remnant vegetation included dense stands of E tereticornis and E crebra

Variable from uniform sands on flats to clay loam duplex soils on slopes



Most of the site was previously used for grazing pasture which was dominated by native grasses and couch Remnant vegetation was dominated by E camaldulensis in the creek lines Paddock trees included E tereticornis E crebra and E acmenoides



E

S

N

W

Figure 4-1 Representation of the localities of the two plantation groups occurring near Bundaberg and Brisbane (black circles)

Northern Plantation Group

Southern Plantation Group

140

Group - Age

Size (ha)


Southern - 4 27914 Undulating with steep crests and deep creek lines

At least half of the site was previously used for grazing pasture which was dominated by native grasses and couch Remnant vegetation was dominated by E camaldulensis in the creek lines Paddock trees included E tereticornis E crebra and E acmenoides

Variable from uniform sands on flats to clay loam duplex soils on slopes (variable depth to saprolite beneath)


Table 42 Site characteristics of the northern plantation group Group - Age Size

(ha) Topography Remnant Vegetation Soil Type Clearing and

Fertiliser History

Northern - 1 4071 Slightly undulating with gradual slopes and shallow creek lines

Most of the site was previously used for grazing pasture which was dominated by native grasses and couch Individual paddock trees included E acmenoides and E crebra Remnant vegetation mainly occurred in blocks along creek lines The dominant species were E tereticornis E intermedia and E exserta

Variable from moderately well drained uniform sandy soils on the flats to granite based and metamorphic sandy loam duplex soils on the slopes



Most of the site was previously used for grazing pasture which was dominated by native grasses and couch Remnant vegetation mainly occurred in blocks along creek lines The dominant species were E tereticornis E intermedia and E exserta

Variable from moderately well drained uniform sandy soils on the flats to granite based sandy loam duplex soils on the slopes Rocky outcrops occur in some areas

Progressive clearing since settlement More recently cleared areas had some regeneration Fertiliser history has been inconsistent


Most of the site was previously used for grazing pasture which was dominated by native grasses couch and rushes in the moister lowland areas Remnant vegetation mainly occurred in blocks along creek lines The dominant species were E tereticornis E intermedia and E exserta

Variable from moderately well drained uniform sandy soils on the flats to granite based sandy clay loam duplex soils on the slopes


Northern - 4 2435 Undulating with steep crests and deep creek lines


Sandy loam duplex soil with medium B-horizons (low salinity)


141

Identifying and Categorising Damage

A preliminary survey of each plantation was conducted to identify the most abundant

pests and pathogens Samples of infected foliage were collected by hand and placed in

paper bags and refrigerated until further examination Insect specimens were stored in

70 ethanol (as described in Chapter 2)

The relative abundance of each form of damage was subjectively estimated at the time

of collection and recorded as high (greater than 60) moderate (between 30-60) or

low (less than 30) Specimens were examined in detail in the laboratory to identify

insects and fungi to species and genus level (Chapters 2 amp 3)

Each pest and pathogen species was placed within a defining causal category (damage

category) Species were placed in damage categories based on the similarity of their

symptoms in plantations These categories also contained taxonomic groupings For

example all damage caused by Teratosphaeria species was allocated to a single

category lsquoTeratosphaeria Damagersquo

As the study progressed new categories were created to include new forms of damage

which were not encountered earlier in the survey Fifteen damage categories were

defined (Table 43)

142

Table 43 A list of the 15 defining damage categories with descriptions of symptoms and causal agents

Damage Category Description of Symptoms Causal Agents Symptoms

Foliar Yellowing

A change in the colour of foliage from green to yellow The incidence may range from a single leaf to the whole canopy The severity may range from minor yellowing such as slight interveinal chlorosis to major yellowing of the entire leaf on both sides Arrows point to yellowing foliage

May have several direct and indirect causes such as A deficiency of water A deficiency of nutrients Damage to leaves and roots by insect pests (causing stress) Damage to host roots by fungal pathogens (cankers) causing stress

Foliar Reddening

A change in the colour of foliage from green to red The incidence may range from a single leaf to the whole canopy being affected The severity may range from minor yellowing such as slight interveinal reddening to major reddening of the entire leaf on both sides Arrow points to red speckling

Caused by the production of anthocyanins in leaf tissues A symptom of stress which may have several direct and indirect causes such as A deficiency of nutrients Damage to leaves and roots by insects pests A change in the colour of foliage from green to yellow (most notably by Psyllids)

143


Physiological Necrosis

The incidence may range from a single leaf to the whole tree being affected The severity may range from minor necrosis such as small patches to entire necrosis of the leaf lamina on both sides Arrow points to necrotic foliage

May have several direct and indirect causes such as A deficiency of water such as a lack of rainfall Stress resulting from damage to roots by insect pests causing moisture stress Damage to host roots by fungal pathogens (cankers) causing moisture stress

Total Fungal Damage

The incidence may range from a single leaf to the whole tree being affected The severity may range from minor necrosis such as small necrotic patches (leaf spots) to entire necrosis of the leaf on both sides (blighting) Different fungal species have different symptoms such as different size and shape and colour of the lesions and different fruiting bodies All fungal pathogens were included in this damage category Arrow points to a necrotic fungal lesion

May be caused by a range of foliar pathogens (see chapter 3)

144


Teratosphaeria Damage

The incidence may range from a single leaf to the whole tree being affected The severity may range from minor necrosis such as small necrotic patches (leaf spots) to entire necrosis of the leaf on both sides Teratosphaeria species can be tentatively identified

in the field by the general appearance of their lesions Lesions usually have defined margins that may be dark brown or red the interior of the lesions are usually light brown to grey and scattered with tiny black fruiting bodies These characteristics were used to identify species in the field which was supported by microscopic examination of samples in the laboratory Arrow points to a Teratosphaeria lesion

May be caused by a range of Teratosphaeria species including M heimii T cryptic T nubilosa M marksii M lateralis

Total Insect Necrosis

The Incidence may range from a single leaf to the whole tree being affected by total insect necrosis The severity may range from minor necrosis such as small necrotic patches to entire necrosis of the leaf on both sides The main difference in distinguishing insect necrosis from physiological necrosis or fungal necrosis is that insect necrosis is usually associated with slight chewing or piercing of the leaf lamina by the mouthparts of the feeding insect All forms of necrosis caused by insect species were included in this category of damage Arrow points to a necrotic lesion

May be caused by a range of insect species which partially consume upper or lower tissues or fluid from the leaf lamina including flea beetles (Galerucinae) amp sap-sucking bugs (Hemiptera)

145


Phylacteophaga Blisters

The incidence may range from a single leaf to the whole tree being affected by Phylacteophaga blisters The severity may range from minor damage such as a few small blisters on the leaf to the entire the leaf being covered in blisters Symptoms of infestation by Phylacteophaga resemble blistering of the leaf surface The adult insects lay eggs within the leaf lamina and the resulting larvae feed on the tissues beneath the cuticle This causes the formation of a blister like structure Arrow points to a leaf blister

May be caused by two species Phylacteophaga froggatti Phylacteophaga eucalypti

Mirid Damage

The incidence may range from a single leaf to the whole tree being affected by Mirid damage The severity may range from minor damage such as a few small necrotic speckles on the leaf to the entire the leaf becoming necrosis Symptoms include feeding scars on the leaf lamina caused by piercing mouthparts and necrotic speckling of the leaf The speckles in low abundance are limited by leaf veins while those in higher abundance usually aggregate into patches Arrow points to necrotic speckling

Caused by Rayiera sp

146


Psyllid Damage

The incidence may range from a single leaf to the whole tree being affected The severity may range from minor damage such as a few lerps (ie protective covering produced by insects) on the leaf to the entire leaf being covered Damage to the leaf is caused by the removal of fluids by the sap-sucking insect beneath the lerp This is often associated with reddening of the tissue around the damaged area Arrows point to lerps on the leaf

Caused by several species including Cardiaspina sp Creiis sp Eucalyptolymma sp

Total Insect Defoliation

The incidence may range from a single leaf to the whole tree being affected by insect defoliation The severity may range from minor damage such a small area of leaf being removed by insect chew to the entire leaf being removed Different defoliating insect species cause different forms of damage The most common method of feeding employed by defoliating insects is chewing the leaf by the mandibles (eg chrysomelid species chew the leaf margins) All forms of insect defoliation were included in this damage category Arrow points to a chewed section of foliage

Caused by several insect species including Chrysomelidae Paropsis atomaria Paropsis obsolete Paropsis variolosa Paropsisterna cloelia Paropsisterna agricola Longitarsus spp Paropsisterna spp Cryptocephalus spp Curculionidae Gonipterus spp Oxyops spp Lepidoptera numerous unidentified species

147


Chrysomelid Defoliation

The incidence may range from a single leaf to the whole tree being affected by Chrysomelid defoliation The severity may range from minor damage such a small area of leaf being removed by chewing to the entire leaf being removed The symptoms of damage by most chrysomelid species are similar and involve scalping of the leaf margin Arrow points to a chewed section of foliage

Many different chrysomelid species including Paropsis atomaria Paropsis obsolete Paropsis variolosa Paropsisterna cloelia Paropsisterna agricola Longitarsus sp Paropsisterna sp Cryptocephalus sp

Weevil Defoliation

The incidence may range from a single leaf to the whole tree being affected The severity may range from minor damage such a small area of leaf being removed by chewing to the entire leaf being removed Symptoms look like a shot gun blast to the foliage of the affected tree each leaf having a series of small circular to irregular holes Larvae tend to feed more voraciously than adults and often consume the entire leaf Slime produced by the larvae may coat the surface of leaves and stems Arrow points to a chewed section of foliage

Caused by species in the genus Gonipterus and Oxyops

Some damage may have accidentally been included which was caused by other Curculionid genera such as Oxyops

148


Foliar Wasp Galls

The incidence may range from a single gall occurring on a single leaf to the whole tree being infested The severity may range from minor damage such as a small gall occurring on the leaf to the entire leaf being occupied by a gall cluster In severe cases branches may snap from the weight of large gall clusters Arrow points to a wasp gall

Caused by several species of wasp in the Chalcidoidea superfamily

Scale Insect Damage

The incidence may range from a single stem to several stems being affected The severity may range from a single scale insect on a stem to several stems being entirely covered in scale insects Symptoms are evident by the presence of scale insects on the stem of the host These resemble aggregations of brown beads and usually affect the lower branches The white coloured individuals are males while the brown individuals are females These usually form separate colonies Sugary secretions produced by the insects often accumulate on foliage and stems near infestations The secretions often become infected with non pathogenic fungi These may cause damage by reducing the photosynthetic area of the leaf Arrow points to a colony of scale insects

Caused by one species Eriococcus coriaceus

149


Leafroller Caterpillars

The incidence may range from a few leaves bound together (occupied by one individual larva) to several such bound structures occupied by several larvae The severity may range from part of a leaf being bound but not chewed by the larva to the whole leaf being consumed by the larva Leaves are bound together with silk by the larva Faecal pellets are also usually associated Arrow points to a cluster of brown necrotic leaves (nest of a leafroller caterpillar)

Caused by the larvae of an unidentified species (Tortricidae)

150

Pest and Disease Assessment Plots

Pest and Disease Assessment Plots (PDA Plots) were established in each plantation

using a method derived from the Crown Damage Index Assessment (CDIA) (Stone et

al 2003) This method involved dividing a map of each plantation into eight equally

sized compartments and then randomly selecting a point within each compartment To

ensure that the points were selected at random a black marker pen was dropped from

head height onto a map lying on a bench by a person with their eyes closed This was

done until a point was selected in each compartment Each point then represented a

location in the plantation at which a PDA plot was established By dividing the plantation

into eight compartments this ensured that assessments occurred throughout the area

of each plantation

Once in the field each of the eight PDA plots were located and marked using a global

positioning system (Magellan GPS Blazer l2) Each plot consisted of a diagonal row of

ten trees (Figure 4-2) Each tree was assessed for pest and disease impacts for a one

hour period A total of 80 trees were assessed within each plantation to give an overall

health status of the plantation at each sampling time

151

Assessing the Incidence and Severity of Damage

The incidence and severity of each damage category was assessed using a modified

version of the Crown Damage Index Assessment (CDIA) by Stone et al (2003) Like the

CDIA the rating system involved estimating two separate measures of damage

lsquoincidencersquo and lsquoseverityrsquo

lsquoIncidencersquo is an estimate of the percentage of the whole tree canopy affected by a

damage category lsquoSeverityrsquo is an estimate of the percentage of damage occurring on

the average leaf

Values were recorded as percentages and rounded to the following measures 5

25 50 75 and 100 The lsquoIncidencersquo and lsquoSeverityrsquo values were then combined

using the following formula to produce a lsquo Total Damagersquo

Total Damage = ( Severity100) times Incidence

Figure 4-2 A diagram representing the structure of a PDA plot within a plantation The green dots represent plantation trees and the hollow dots represent trees included in the assessment

152

When assessing the Total Damage for each damage category this system was

applied to each tree within the PDA plot which was then averaged (ten trees)


The abundance of pests and pathogens in different aged plantations was compared by

comparing the Total Damage for each damage category between different aged

plantations Age comparisons were made within both the northern and southern

plantation groups


The climatic characteristics of the northern and southern plantation groups were

identified using long term weather data from the Australian Bureau of Meteorology

(wwwbomgovau) The Amberley Weather Station (station 040004) supplied data

(within 25 km) for the southern plantation group and the Town of 1770 Weather Station

(station 039314) provided data for the northern plantation group (within 10 km)

The Total Damage was compared between the northern and southern plantation

groups for each damage category (equally aged plantations) (Table 44)

Table 44 Paired comparisons of equal aged plantations in the northern and southern plantation groups

Plantation (Group ndash Age)

North-1 South-1 North-2 South-2 North-3 South-3 North-4 South-4


The PDA plots were assessed at three month intervals during a twelve month period

(August 2004 November 2004 February 2005 and May 2005) BOM data were used to

correlate weather patterns with the seasonal abundance of pests and pathogens

153

Statistics and Multivariate Analyses

All data were collected in the field using a portable palmtop computer (HP Pavilion)

Data were entered into an Excel data spreadsheet during each site visit (Microsoft)

Multivariate analyses were carried out using the Primer 5 statistical package The Bray-

Curtis similarity coefficient was employed to construct a similarity matrix from the log

(n+1) transformed values of each damage category This matrix was then subjected to

non-metric multidimensional scaling (MDS) ordination One way crossed Analysis of

Similarities (ANOSIM) was carried out to ascertain whether the compositions of the

damage categories differed significantly between four different aged plantations

northern versus southern plantation groups and four different seasons The factors

employed in each of the tests are specified in the results In each test the null

hypothesis lsquothat there were no significant differences among groupsrsquo was rejected if the

significance level (P) was lt5 The R statistic value was used to ascertain the extent of

any significant differences Any R values lt01 were regarded as negligible Where

ANOSIM detected a significant difference among priori groups and the R-statistic was

gt01 similarity percentages (SIMPER) were used to identify which damage categories

made the greatest contribution to those differences

Results

Damage Averages

A comparison of the Total Damage for each damage category showed that most

damage was caused by insect groups (Table 45) Total Defoliation caused the highest

Total Damage (averaged across all plantations) Since most of the damage within

this category was caused by chrysomelid beetles it is not surprising that the second

highest measure of damage was caused by Chrysomelid Damage Other high

154

measures of damage included Total Insect Necrosis Physiological Necrosis and Foliar

Yellowing All other damage categories caused 41 or less of the total damage

recorded

Damage category Total Damage Rank (High-Low)

Total Insect Defoliation 295 1st

Chrysomelid Damage 265 2nd

Total Insect Necrosis 119 3rd

Physiological Necrosis 82 4th

Foliar Yellowing 77 5th

Foliar Reddening 41 6th

Total Fungal Damage 30 7th

Teratosphaeria Leaf Blight 29 8th

Mirid Damage 29 9th

Leafroller Caterpillars 17 10th

Foliar Wasp Galls 07 11th

Phylacteophaga Blisters 06 12th

Weevil Defoliation 01 13th

Scale Insect Damage 01 14th

Psyllid Damage 01 15th

Total 100

A comparison of the average Total Damage (all damage categories) between

different aged plantations showed that one-year-old plantations had the lowest levels of

damage followed by three-year-olds four-year-olds and two-year-olds (Table 46) The

northern plantation group had a higher average Total Damage than the southern

plantation group A comparison of the average Total Damage between seasons

showed that the highest levels of damage occurred in May 2005 followed by February

2005 August 2004 and November 2004 Higher levels of damage occurred in the

second half of the study period

Table 45 Average Total Damage (all categories) for each damage category

155

Plantation Age

One-Year-Old Two-Year-Old Three-Year-Old Four-Year-Old

34 60 43 53

Local Climate

Southern Plantation Group Northern Plantation Group

37 58

Seasons

August 2004 November 2004 February 2005 May 2005

41 30 59 60


Multivariate statistics were used to collectively compare damage category data between

the different aged plantations A one way crossed analysis of similarities showed that

collective levels of damage varied significantly (Plt05 Rgt01) between different aged

plantations within each plantation group (Table 47 Table 48)

The Global R value of the southern plantation group (0346) was less than the Global R

value of the northern plantation group (0580) which infers that that there were greater

differences (more variability) in collective measures of damage between plantations in

the northern plantation group

Month (P=01 Global R=0346)

Southern Plantation Group (aged 1-4 yrs)

Southern Plantation Group (1-4 years)

South-1 South-2 South-3 South-4

P R P R P R P R South-1 South-2 01 0669 South-3 01 0343 01 0107

South-4 01 0660 01 0231 01 0179

(P=01 Global R=0580)

Northern Plantation Group (aged 1-4 yrs) Northern Plantation Group (1-4 years)

North-1 North-2 North-3 North-4

P R P R P R P R North-1 North-2 01 0915 North-3 01 0898 01 0215 North-4 01 0896 01 0382 01 0226

Table 48 Significance levels P and R statistic values for both global and pair-wise comparisons in a one way ANOSIM test of all measures of damage in the Northern plantations (North-1 North-2 North-3 North-4) respectfully Significant results (Plt05 Rgt01)

Table 47 Significance levels P and R statistic values for both global and pair wise comparisons in a one way ANOSIM test of all measures of damage in the Southern Plantation Group (South-1 South-2 South-3 amp South-4) respectfully Significant results (Plt05 Rgt01)

Table 46 Average Total Damage (all categories) for Plantation Age Local Climate and Season

156

Multi dimensional scaling using ordination was used to compare collective measures of

damage between different aged plantations between the northern and southern

plantation groups (Figure 4-3) The analysis showed a distinct separation by distance of

the points representing the one-year-old southern and northern plantations from the

other differently aged plantations The stress value being lt2 (The degree of

correspondence between the distances among points) implied that the MDS map and

matrix input displayed an ordination that was an acceptable representation of the

observed variability in the analysis The ordination supported what was suggested by

ANOSIM (Table 47 Table 48) that the 1-year-old plantations in the southern and

northern plantation groups were most dissimilar in terms of collective measures of

damage The ordination also showed that the one-year-old plantations in the northern

and southern groups were similar to each other

North-3

Figure 4-3 Two dimensional MDS ordination of the second-stage similarity matrix containing the correlations between each pair of similarity matrices constructed from measures of damage for all damage categories for all samples in the southern plantations (south-1 south-2 south-3 amp south-4) and the northern plantations (north-1 north-2 north-3 north-4) Each point can be identified by its corresponding plantation

Stress 017 Group of one-year -old plantations

South-1

South-2

South-3

South-4

North-1

North-2

North-4

North-3

157

Similarity percentages (SIMPER) (Clarke and Gorley 2001) were used to identify which

damage categories made the greatest contribution to differences between plantations in

terms of collective measures of damage Total Insect Defoliation Chrysomelid

Defoliation and Total Insect Necrosis were ranked as the greatest contributors in all four

plantations within the southern plantation group (Table 49) These damage categories

were also amongst the greatest contributors in plantations within the northern plantation

group with the exception of Foliar Reddening which was the greatest contributor in the

one-year-old northern plantation (Table 410) This was expected given that these

damage categories generally caused the greatest Total Damage

158

Southern Plantation Group (aged 1-4 years)

Rank South-1 South-2 South-3 South-4

1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th

13th

Chrysomelid Defoliation (81) Total Insect Defoliation (87) Total Insect Necrosis (48)

Phylacteophaga Blisters (04) Foliar Wasp Galls (00) Foliar Yellowing (32) Total Fungal Damage (04) Physiological Necrosis (116) Teratosphaeria Damage (04) Mirid Damage (03)

Total Insect Defoliation (276) Chrysomelid Defoliation (202) Total Insect Necrosis (63)

Total Fungal Damage (12) Teratosphaeria Damage (12) Foliar Yellowing (31) Physiological Necrosis (130) Phylacteophaga Blisters (00)

Total Insect Necrosis (55) Total Insect Defoliation (276) Chrysomelid Defoliation (113)

Total Fungal Damage (04) Foliar Yellowing (38) Teratosphaeria Damage (04) Physiological Necrosis (79) Phylacteophaga Blisters (00) Foliar Wasp Galls (00) Mirid Damage (00) Foliar Reddening (00) Eucalypt Leafroller Caterpillar (04) Scale Insect Damage (00)


Total Fungal Damage (02) Foliar Yellowing (40) Physiological Necrosis (116) Teratosphaeria Damage (01) Phylacteophaga Blisters (00) Foliar Wasp Galls (00) Mirid Damage (00)

Northern Plantations (aged 1-4 years)

Rank North-1 North-2 North-3 North-4

1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th

13th

14th

15th

Foliar Reddening (95) Total Insect Necrosis (81) Total Insect Defoliation (131) Chrysomelid Defoliation (128)

Eucalypt Leafroller Caterpillars (53) Teratosphaeria Damage (15) Total Fungal Damage (15) Phylacteophaga Blisters (03) Foliar Wasp Galls (11) Mirid Damage (39) Foliar Yellowing (13) Physiological Necrosis (04) Scale Insect Damage (00) Psyllid Damage (01) Weevil Defoliation (00)


Total Fungal Damage (71) Teratosphaeria Damage (69) Foliar Yellowing (69) Mirid Damage (39) Foliar Reddening (08) Physiological Necrosis (08)

Total Defoliation (216) Chrysomelid Defoliation (215) Total Insect Necrosis (89)

Mirid Damage (37) Total Fungal Damage (35) Teratosphaeria Damage (35) Foliar Yellowing (21) Physiological Necrosis (05) Phylacteophaga Blisters (12) Foliar Wasp Galls (00)


Total Fungal Damage (36) Teratosphaeria Damage (35) Foliar Yellowing (21) Mirid Damage (10) Phylacteophaga Blisters (10)

Table 49 Damage categories detected by SIMPER as having the greatest contribution to the differences in collective measured damage between the Southern plantations (South-1 South2 South-3 amp South-4) Ranked from greatest to lowest contribution (bold categories are considered to be the major contributers to collective damage) Average percentage () of damage included in brackets

Table 410 Damage categories detected by SIMPER as having the greatest contribution to the differences in collective measured damage between the Northern plantation group (North-1 North-2 North-3 North-4) Ranked from highest to lowest contribution (bold categories are considered to be the major contributers to collective damage) Average percentage () of damage included in brackets

159

Measures of damage varied greatly between plantations for almost all damage

categories included in the study (Figure 4-7 Figure 4-8 Figure 4-9 Figure 4-10 Figure

4-11 Figure 4-12 Figure 4-13 Figure 4-14 and Figure 4-15) Damage categories which

did not show large variability were generally those which occurred in low abundance

These included Eucalypt Leafroller Caterpillars Foliar Wasp Galls Phylacteophaga

Blisters Weevil Defoliation Scale Insect Damage and Psyllid Damage (Table 411 and

Table 412)

160

Total Insect Chrysomelid Total Insect Physiological Foliar Foliar Total FungalMycosphaerella Mirid Leafroller Foliar WaspPhylacteophaga Weevil Scale Insect Psyllid

Defoliation Defoliation Necrosis Necrosis Yellow ing Reddening Damage Damage Damage Caterplillar Galls Blisters Defoliation Damage Damage

mean 23 18 1 34 14

plusmn SE 14 1 07 44 19

mean 3 3 13 19 16 16 01

plusmn SE 13 13 08 37 13 13 02

mean 19 16 08 28 04 04 01

plusmnSE 16 11 07 45 09 09 02

mean 19 16 08 28 04 04 01

plusmnSE 08 08 1 46 02

mean 22 2 12 27 05 05 04

plusmnSE 13 12 09 42 1 1 11 01

mean 19 16 08 28 04 04

plusmn SE 1 09 09 15

mean 184 183 116 28 28

plusmn SE 6 59 58 41 41

mean 121 12 121 125 04 04

plusmn SE 57 57 67 354 06 06

mean 145 13 25 13 03

plusmn SE 29 21 46 35 05

mean 116 111 116 37 03 08 08 01

Southern plusmnSE 75 74 81 173 17 23 23 03

Plantation mean 152 138 152 43 06 06 13 09 24

Group plusmn SE 79 79 66 37 12 12 23 2 4

mean 519 222 519 33 02 09 3 03

plusmn SE 74 74 68 7 05 27 69 05

mean 191 143 191 53 01 01 16 04

plusmnSE 116 68 116 45 02 02 26 07

mean 421 155 421 56 02

plusmnSE 75 75 53 72 04

mean 321 165 321 46 02 02 04 09 07 06 02

plusmnSE 173 78 173 56 06 06 12 21 35 22 04

mean 152 152 175 313 22 05 05

plusmnSE 45 45 32 17 87 06 06

mean 371 371 124 7 02 02 88 13

plusmn SE 72 72 25 35 04 04 12 35

mean 174 174 9 71 05 05

plusmn SE 58 58 17 14 07 06

mean 245 245 113 41 77

plusmn SE 85 85 41 82 83

mean 235 235 126 88 15 03 03 22 03

plusmnSE 18 18 43 16 96 05 05 69 18

Minor Damage categoriesMajor Damage Categories

Nov-04

1

2

1

2

3

4

4

Total

1

Percentage of Damage ()

Total

3

4

Total

1

2

3

2

3

4

Total

AgeEstate month

May-05

Feb-05

Aug-04

Table 411 A summary of percentage means plusmn SE for each damage category in the southern plantation group (1 2 3 and 4 years old) in August 2004 November 2004 February 2005 and May 2005

161



mean 33 13 26 38 378 01

plusmn SE 28 28 25 16 165 04

mean 16 154 141 163 32 179 177 145 19 01

plusmn SE 52 48 46 85 17 5 5 46 37 04

mean 135 13 155 222 141 93 92 146 16 01 05

plusmn SE 68 68 52 179 15 4 4 37 35 02 08

mean 343 341 145 43 94 67 65 38 39

plusmn SE 64 64 45 137 18 48 48 5 4

mean 168 159 117 26 229 85 83 83 18 01

plusmnSE 13 13 67 183 185 76 75 75 34 02 05

mean 9 9 55

plusmn SE 55 55 09

mean 9 9 15 44 24 23 147

plusmn SE 38 38 72 73 24 24 69

mean 29 29 75 25 3 3 02 13

plusmn SE 51 51 1 46 22 22 07 11

mean 172 171 92 29 67 67 01

plusmn SE 22 22 44 76 39 39 02

Northern mean 139 139 93 24 29 29 38 03

Plantation plusmnSE 68 68 54 56 33 33 73 08

Group mean 238 238 5 06 58 58 11 163 06

plusmn SE 25 2 18 72 72 18 132 18

mean 35 35 15 63 78 77 26 01

plusmn SE 113 113 34 92 57 55 4 02

mean 34 34 111 17 17 16 14 02 01

plusmn SE 125 125 43 29 09 09 24 04 02

mean 356 356 8 13 05 05 03

plusmn SE 57 57 2 23 07 07 07

mean 321 321 86 25 39 39 03 47 02 04 01

plusmnSE 99 98 37 53 53 53 1 95 09 13 04 02

mean 17 17 193 16 09 03 145 47 38 12 03

plusmn SE 43 43 56 31 27 07 8 14 58 2 09

mean 443 443 12 3 04 04 15 04

plusmn SE 33 33 2 3 09 05 11 04

mean 36 36 16 2 16 01 13 03

plusmn SE 62 62 17 28 35 03 19 09

mean 464 464 13 16 03 02

plusmn SE 87 87 27 23 04 04

mean 359 359 133 21 07 03 36 41 09 04 02

plusmnSE 131 131 49 27 22 05 02 74 83 32 11 05

Minor Damage CategoriesMajor Damage Categories

Total

Total

2

Nov-04

1

2

1

2

3

4

3

4

1

2

3

4

Total

1

3

4

Total


month AgeEstate

May-05

Aug-04

Feb-05

Table 412 A summary of percentage means plusmn SE for each damage category in the northern plantation group (1 2 3 and 4 years old) in August 2004 November 2004 February 2005 and May 2005

162


Multivariate statistics were used to collectively compare damage levels between the

northern and southern plantation groups To reduce the effects of confounding variables

only the equally aged plantations were compared in the analysis (Table 413) A one-

way crossed analysis of similarities (ANOSIM) showed that significant (Plt05 Rgt01)

differences occurred when comparing the one-year-old northern and southern

plantations the two-year-old northern and southern plantations and the four-year-old

northern and southern plantations (Table 413) Based on this analysis the null

hypothesis that there were no significant differences between the plantation groups is

rejected for the one two and four-year-old plantations



P R P R P R P R North-1 01 0153 North-2 03 0122 North-3 37 004 North-4 01 0136

Multi-dimensional scaling using ordination was used to compare collective measures of

damage between the two plantation groups This analysis showed very little separation

of the points representing plantations within each plantation group (Figure 4-4) The

stress value (lt2) indicated that the ordination was an acceptable representation of the

observed variability between measurements within the analysis

Table 413 Significance levels (p) and R ndashstatistic values for both global and pair-wise comparisons in a one way ANOSIM test of all damage categories between plantations of the same age in the different plantation groups (Southern Plantation Group South-1 South2 South-3 amp South-4 Northern Plantation Group North-1 North-2 North-3 amp North-4) Significant results (Plt05 Rgt01)

163

When comparing R values from the previous one way crossed analysis of similarities

within plantation groups (Table 47 Table 48) to the one way crossed analysis between

plantation groups (Table 413) the differences between plantations within each

plantation group appears to be more significant than the differences between the

plantation groups This is especially true for the one-year-old plantations (north and

south) which suggests that these plantations have greater similarity than the

plantations within their corresponding groups This is also supported by the grouping

displayed in the previous ordination (Figure 4-3)


damage categories made the greatest contribution to differences between the northern

and southern plantation groups Total Defoliation Chrysomelid Defoliation and Total

Insect Necrosis were ranked as the highest contributors in both plantation estates

(Table 414) This was expected given that these damage categories caused the

greatest Total Damage (Table 45)

Figure 4-4 Two dimensional MDS ordination of the second-stage similarity matrix containing the correlations between each pair of similarity matrices constructed from the severity of damage categories in all samples (Southern and Northern Plantations) Each point can be identified by its corresponding plantation estate

Stress 017 No groupings

Southern Plantations

Northern Plantations

164

Plantations Estates

Rank Southern Plantations Northern Plantations

1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th


Physiological Necrosis (110) Total Fungal Damage (05) Foliar Yellowing (35) Mirid Damage (01) Teratosphaeria Damage (05) Foliar Wasp Galls (07) Eucalypt Leafroller Caterpillars (02) Phylacteophaga (01)

Total Defoliation (236) Chrysomelid Defoliation (234) Total insect Necrosis (90)

Teratosphaeria Damage (38) Foliar Yellowing (43) Mirid Damage (40) Total Fungal Damage (39) Foliar Reddening (40) Foliar Yellowing (43) Foliar wasp Galls (03) Phylacteophaga Blisters (07) Eucalypt Leafroller Caterpillars (17)

Climate Averages

Long term temperature data (1941-2008) showed a year long trend of higher mean daily

maximum temperature in the southern plantation group compared with the northern

plantation group (Figure 4-5A) However mean daily minimum temperature was higher

in the northern plantation group (Figure 4-55B) This indicates that overall the northern

plantation group has a warmer climate during most stages of the year Typical

temperatures were experienced in both plantation groups during the study period

(Figure 4-6C amp D)

Long term rainfall data (1941-2008) shows that the northern and southern plantation

groups typically received low rainfall from July-September and high rainfall from

December-February (Figure 4-5A amp B) Annual rainfall is also typically greater in the

northern plantation group However during the study period both plantation groups

experienced extremely dry conditions from July ndash September 2004 and only the

northern plantation group received normal rainfall from December 2004 ndash February

2005 (Figure 4-5C amp D)

Table 414 Damage categories detected by SIMPER as having the greatest contribution to the differences in collective measured damage between the southern and northern plantation estates Ranked from highest to lowest contribution (bold categories are considered to be the major contributers to collective damage) Average percentage () of damage included in brackets

165

Observations in the field showed that when high rainfall was received in plantations

they responded by producing copious amounts of new foliage (flush growth) Prolonged

periods without rain caused moisture stress which led to high Physiological Necrosis

Foliar Yellowing and leaf loss By the end of the study period areas in which the

northern and southern plantation groups occurred were declared to be severely drought

stricken (Queensland Drought Report May 2005)

May Jun July Aug Sep Oct Nov Dec Jan Feb Mar Apr May May Jun July Aug Sep Oct Nov Dec Jan Feb Mar Apr May

0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

Mean Rainfall (mm)

Mean Max Temp (C)

Mean Min Temp (C)


Months Months

Figure 4-5 A Amberley weather station data 1941-2008 B Town of 1770 weather station data 1941-2008 C Amberley weather station data 2004-2005 D Town of 1770 weather station data 2004-2005 Mean maximum daily temperature () mean minimum daily temperature () and mean monthly rainfall (prod) Australian Bureau of Meteorology

0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300M

ean

Rain

fall (

mm

)

Mean Rainfall (mm)

Mean Max Temp (C)

Mean Min Temp (C)

START OF SURVEY

C D

2004 2005 2005 2004

A B

166



seasons A one way crossed analysis of similarities (ANOSIM) showed that collective

levels of damage varied significantly (Plt05) between all four seasons of sampling

(Table 415) R values from this analysis indicated that the most different season in

terms of collective measures of damage was May 2005 which was most dissimilar to

November 2004 and August 2004


Aug 04 Nov 04 Feb 05 May 05

P R P R P R P R

Aug 04

Nov 04 01 0438

Feb 05 01 0631 01 0547

May 05 01 091 01 0934 01 077

Multi Dimensional Scaling (MDS) using ordination (ie dissimilarity by distance) was

used to compare collective measures of damage between seasons The MDS showed a

distinct separation by distance of the points representing collective measures of

damage for August 2004 and May 2005 (Figure 4-6) The stress value (lt2) indicated

that the ordination was an acceptable representation of the observed variability between

the measurements in the analysis The ordination was consistent with what was

suggested by ANOSIM that May 2005 was the most different season followed by

August 2004 November 2004 and February 2005 (Table 415) Greater separation by

distance was observed for the seasonal ordination than previous analyses This may

also suggest that season has a greater influence on collective measures of damage

than both plantation age and local climate

Table 415 Significance levels (p) and R ndashstatistic values for both global and pair wise comparisons in a one way ANOSIM test of all damage categories across all plantations in all four seasons of sampling (Significant results (Plt05 Rgt01))

167


damage categories made the greatest contribution to differences between seasons in

terms of collective measures of damage Total Defoliation Chrysomelid Defoliation and

Total Insect Necrosis were ranked among the highest contributors in August 2004

November 2004 and February 2005 (Table 416)

Figure 4-6 Two dimensional MDS ordination of the second-stage similarity matrix containing the correlations between each pair of similarity matrices constructed from the severity of all fifteen damage category all samples (eight plantations four seasons) The points are coded for season The analysis contains four groups

Four Groupings

Stress 017

168

Rank

Seasons


1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th


Total Fungal Damage (45) Foliar Yellowing (72) Teratosphaeria Damage (45) Mirid Damage (41) Foliar Reddening (81) Phylacteophaga Blisters (11) Foliar Wasp Galls (00) Physiological Necrosis (00) Eucalypt Leafroller Caterpillar (00)

Chrysomelid Defoliation (104) Total Defoliation (106) Total Insect Necrosis (80)

Teratosphaeria Damage (20) Mirid Damage (19) Phylacteophaga Blisters (02) Total Fungal Damage (20) Physiological Necrosis (16) Foliar Wasp Galls (00) Foliar Yellowing (16) Foliar Reddening (02) Eucalypt Leafroller Caterpillar (00)


Teratosphaeria Damage (20) Total Fungal Damage (21) Eucalypt Leafroller Caterpillar (28) Foliar Yellowing (36) Mirid Damage (03) Physiological Necrosis (160) Phylacteophaga Blisters (02) Foliar Wasp Galls (05)

Total Defoliation (297) Chrysomelid Defoliation (297) Physiological Necrosis (55)

Teratosphaeria Damage (02) Mirid Damage (18) Foliar Gall Wasps (16) Foliar Reddening (00) Total Insect Necrosis (107) Total Fungal Damage (03)

Table 416 Damage categories detected by SIMPER as having the greatest contribution to the differences in collective measured damage between the seasons (all plantations) Ranked from highest to lowest contribution (bold categories are considered to be the major contributers to collective damage) Mean percentage () of damage included in brackets

169

Total Insect Defoliation and Chrysomelid Defoliation

Total Insect Defoliation and Chrysomelid Defoliation showed very similar patterns of

abundance throughout the study period because chrysomelid beetles caused over 90

of the damage contributing to Total Insect Defoliation Only in February 2005 in the

southern plantation group did Total Insect Defoliation occur at noticeably higher levels

than Chrysomelid Defoliation (Figure 4-7 and Figure 4-8) Other insect groups are likely

to have caused higher levels of damage during this period

The Total Damage for Total Insect Defoliation and Chrysomelid Defoliation was

highly variable within both plantation groups Levels of damage were consistently low in

the one-year-old plantations compared with the two three and four-year-old plantations

Because the Total Damage was highly variable within both plantation groups

differences between the plantation groups were difficult to detect Only one clear

difference between plantation groups occurred in August 2004 when the Total

Damage was consistently lower in the southern plantation group than the northern

plantation group

Seasonal differences in Total Insect Defoliation and Chrysomelid Defoliation were

difficult to detect but higher levels of damage occurred in the second half of the study

period than the first half

170

Figure 4-7 Total Insect Defoliation (plusmn SE) Total Damage for 1 to 4-year-old plantations in four seasons of sampling (August 2004 November 2004 February 2005 and May 2005) in the southern plantation group (A-D) and the northern plantation group (E-H)

00

100

200

300

400

500

600

Bri-1 Bri-2 Bri-3 Bri-4

Plantation (Estate-Age)

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

Bun-1 Bun-2 Bun-3 Bun-4


Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600


Plantation (Estate-Age)P

erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










171

Figure 4-8 Chrysomelid Defoliation Total Damage (plusmn SE) for 1 to 4-year-old plantations in four seasons of sampling (August 2004 November 2004 February 2005 and May 2005) in the southern plantation group (A-D) and the northern plantation group (E-H)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










172


The Total Damage of Total Insect Necrosis was more variable throughout the study

period in the southern plantation group than the northern plantation group (Figure 4-9)

In August 2004 and February 2005 the Total Damage almost disappeared in the

southern plantation group while remaining between 5-15 throughout most of the

study period in the northern plantation group

The Total Damage was consistently lower in the one-year-old plantations of both

plantation groups throughout the study period with the exception of the final

assessment in May 2005 in which levels were highest in the one-year-old plantations

The main difference in the Total Damage between plantation groups was that low

levels were observed in the southern plantation group in August 2004 and February

2005 Seasonal changes appeared to be more prevalent in the southern plantation

group with levels of damage changing more significantly between samples


Physiological Necrosis was absent from the southern plantation group in August 2004

and absent from the northern plantation group until the final sample in May 2005 (Figure

4-10) Given the high levels of damage observed in February 2005 in the southern

plantation group this form of damage was probably the most variable of all damage

categories

No clear patterns of abundance were observed when comparing different aged

plantations with each plantation group The highest level of damage occurred in a three-

year-old plantation in November 2004 a two-year-old plantation in February 2005 and a

one-year-old plantation in 2005 Seasonal effects appear to be strongest in the southern

plantation group with highly variable levels of damage between seasonal samples

173

Figure 4-9 Total Insect Necrosis Total Damage (plusmn SE) for 1 to 4-year-old plantations in four seasons of sampling (August 2004 November 2004 February 2005 and May 2005) in the southern plantation group (A-D) and the northern plantation group (E-H)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










174

Figure 4-10 Physiological Necrosis Total Damage (plusmn SE) for 1 to 4-year-old plantations in four seasons of sampling (August 2004 November 2004 February 2005 and May 2005) in the southern plantation group (A-D) and the northern plantation group (E-H)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










175

Foliar Yellowing and Foliar Reddening

The Total Damage of Foliar Yellowing was low (lt10) in all plantations during the

study period with the exception of the southern plantation group in May 2005 and the

northern plantation group in August 2004 (Figure 4-11) Foliar Reddening only occurred

at very low levels in November 2004 in a four-year-old plantation within the southern

plantation group and at higher levels in all four plantations of the northern plantation

group in August 2004 (Figure 4-12)

Although Foliar Yellowing and Foliar Reddening occurred at low levels during most of

the study period it is interesting that both forms of damage occurred at high levels in

the northern plantation group in August 2004 This may suggest that local climate or

season was having an influence It is also interesting that when comparing different

aged plantations during this time the two forms of damage have opposite patterns of

abundance Foliar Yellowing shows increasing levels of damage with increasing

plantation age while Foliar Reddening shows decreasing levels of damage with

increasing plantation age

176


Figure 4-11 Foliar Yellowing Total Damage (plusmn SE) for 1 to 4-year-old plantations in four seasons of sampling (August 2004 November 2004 February 2005 and May 2005) in the southern plantation group (A-D) and the northern plantation group (E-H)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F August 2004 November 2004 February 2005 May 2005

G H









North-1 North-2 North-3 North-4 North-1 North-2 North-3 North-4 North-1 North-2 North-3 North-4 North-1 North-2 North-3 North-4 North-1 North-2 North-3 North-4

177

Figure 4-12 Foliar Reddening Total Damage (plusmn SE) for 1 to 4-year-old plantations in four seasons of sampling (August 2004 November 2004 February 2005 and May 2005) in the southern plantation group (A-D) and the northern plantation group (E-H)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H











178

Total Fungal Damage and Teratosphaeria Damage

Total Fungal Damage (Figure 4-13) and Teratosphaeria Damage (Figure 4-14) showed

very similar patterns of abundance throughout the study period This is because Total

Fungal Damage contributed to over 90 of the damage within the Total Fungal

Damage category

The greatest levels of Total Damage occurred in a two-year-old and three-year-old

plantation in the northern plantation group in August 2004 Given that levels of damage

were consistently low in other plantations during the study period no patterns of

abundance are apparent when comparing different aged plantations

Levels of damage were consistently higher in the northern plantation group than the

southern plantation group

179

Figure 4-13 Total Fungal Damage Total Damage (plusmn SE) for 1 to 4-year-old plantations in four seasons of sampling (August 2004 November 2004 February 2005 and May 2005) in the southern plantation group (A-D) and the northern plantation group (E-H)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










180



Figure 4-14 Teratosphaeria Damage Total Damage (plusmn SE) for 1 to 4-year-old plantations in four seasons of sampling (August 2004 November 2004 February 2005 and May 2005) in the southern plantation group (A-D) and the northern plantation group (E-H)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H








South-1 South-2 South-3 South-4 South-1 South-2 South-3 South-4


181

Mirid Damage

Mirid Damage was largely absent from the southern plantation group during most of the

study period although low levels occurred in a one-year-old plantation and a two-year-

old plantation in February 2005 (Figure 4-15) Damage was detectable throughout the


Interestingly damage was greater in the two-year-old and three-year-old plantations

during the first half of the study period and then greater again in the one-year-old

plantations during the second half of the study period This makes it difficult to attribute

changes in damage to either plantation age or seasonal effects

Mirid Damage was consistently higher in the northern plantation group than in the

southern plantation group which suggests that mirid damage may be under the

influence of local climate

182

Figure 4-15 Mirid Damage Total Damage (plusmn SE) for 1 to 4-year-old plantations in four seasons of sampling (August 2004 November 2004 February 2005 and May 2005) in the southern plantation group (A-D) and the northern plantation group (E-H)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H









North-1 North-2 North-3 North-4 North-1 North-2 North-3 North-4 North-1 North-2 North-3 North-4 North-1 North-2 North-3 North-4

183

Low Damage Categories

Damage caused by eucalypt leafroller caterpillars foliar wasp galls phylacteophaga

blisters weevil defoliation scale insect damage and psyllid damage collectively caused

only 51 of the total damage in the southern plantation group and 62 of the total

damage in the northern plantation group (Table 411 and Table 412 respectfully) These

damage categories are therefore considered to have negligible impacts

Eucalypt leafroller caterpillars only affected plantations in the second half of the study

period at low levels The highest level of damage recorded was 16 Total Damage which

occurred in a one-year-old plantation in the northern plantation group

Foliar wasp galls also only occurred in the second half of the study period at low levels

The galls appeared to similarly affect different aged plantations in both groups The

highest level of damage recorded was 88 Total Damage which occurred in a two-year-

old plantation in the northern plantation group

Phylacteophaga blisters only caused low levels of damage in the southern plantation

group in August 2004 and was absent in all subsequent seasons of sampling Similarly low

levels of damage affected different aged plantation within the northern plantation group

The highest level of damage recorded was 16 Total Damage which occurred in a four-

year-old plantation in the northern plantation group

Weevil defoliation only occurred at low levels in both plantation groups during February

2005 The highest level of damage recorded was 24 Total Damage which occurred in a

one-year-old plantation in the southern plantation group

Scale insect damage and psyllid damage caused the lowest levels of damage during the

study period and occurred sporadically in both plantation groups at very low levels (mostly

less than 1)

184

Discussion

Drought in Southern Queensland

Atypical climatic conditions occurred in southern Queensland during the study period

Although southern Queensland generally experiences high rainfall and temperatures

during the summer months the region was declared drought stricken in May 2005 due to a

severe lack of rainfall in many areas (Queensland Drought Report May 2005) The impacts

of drought appeared to be greater in the southern plantation group which received less

rain during summer Field observations indicated that rainfall events were often extremely

localised On several occasions plantations were observed receiving rain while nearby

plantations (lt10 km) received no rain This observation illustrated that weather station

data which was collected approximately 10 km from each plantation group could only be

used as a rough guide as to the amount of rain actually received by plantations

Within plantations the processes of leaf loss and regeneration were observed to be

accelerated by drought conditions Because damage was measured using a proportion

based system (percentage of damaged foliage versus healthy foliage) the processes of

leaf loss and subsequent regeneration after rainfall had a confounding effect on the study

Moisture stressed trees tended to lsquodroprsquo foliage which was already damaged by pests and

pathogens Therefore leaf loss could cause a direct decrease in the percentage of

damaged leaves in tree canopies Similarly the production of new healthy foliage after rain

could cause a decrease in the percentage of damaged leaves (dilution effect) In other

words it was difficult to attribute any changes in damage to actual changes in the

population size of pest or pathogens because any change could equally be attributed to

the effects of leaf loss or regeneration

Eucalypts are capable of continuous growth and may recover quickly after damage by fire

herbivore damage or drought (Jacobs 1955 Beadle and Inions 1990) This was also

185

observed in the Queensland plantations and single rainfall events could dramatically

improve the overall health of plantations Other changes in canopy health such as wilting

and senescence appeared to occur more gradually during periods of moisture stress

When the time between rainfall events was prolonged this resulted in high moisture

stress These plantations would go through rapid cycles of leaf loss and subsequent

regeneration after rainfall It was soon realised that these effects had the potential to

overshadow more gradual accumulative effects such as plantation age regional climate

and season


Many insects and pathogens prefer juvenile foliage of eucalypts (Macauley and Fox 1980

Abbott 1993 Day 1998 Larsson and Ohmart 1998 Steinbauer et al 1998 Brennan et al

2001 Lawrence et al 2003) It was therefore expected that the abundance of pests and

pathogens would be greater in young plantations where juvenile foliage was more

abundant Contrary to this expectation the findings of the study revealed that the majority

of damage categories caused low levels of damage in the one-year-old plantations

(observed in both northern and southern plantation groups) Total Insect Defoliation

Chrysomelid Defoliation and Total Insect Necrosis caused the highest levels of damage

during the study period however these levels were lowest in the one-year-old plantations

This may have been attributed to a faster rate of regrowth in these plantations Younger

plantations were also observed to drop their foliage very quickly during periods of high

moistures stress while older plantations tended to resist drought better and retained their

foliage It is therefore likely that younger plantations replaced damaged foliage quicker

than older plantations Such effects could lead to lower measurable damage by insects in

younger plantations despite higher feeding rates of insects

Mirid damage (Rayieria sp) and leafroller caterpillars (Stepsicrates sp) caused higher

186

levels of damage in the two-year-old plantations Both species appeared to have a

preference for soft juvenile leaves rather than tough mature leaves Although one-year-old

plantations may well have been more attractive to these pests both species appeared to

build population numbers slowly This may help explain why greater levels of damage

occurred in two-year-old plantations (gradual build up) A higher proportion of mature

foliage to juvenile foliage in three and four-year-old plantations may also have made these

plantations less attractive to pest species

The effects of drought in southern Queensland largely overshadowed the effects of

plantation age This was mainly because all stressed plantations regardless of age

tended to produce new regrowth This made the canopy characteristics of differently aged

plantations similar It is expected that the drought may have facilitated insect pests by

increasing availability of palatable foliage thereby predisposing trees to greater

infestations


In August 2004 plantations in the southern plantation group were observed to be in a

moderately good state of health Local people reported that very little rain had occurred

over the previous months but most plantation trees appeared to be enduring the dry

conditions In November 2004 most of the plantations had received at least some summer

rainfall which caused them to produce large quantities of new foliage This was

particularly evident in the younger plantations In February 2005 after a drier than average

summer the plantations began to show symptoms of moisture stress which caused large

scale wilting of foliage and leaf loss In May 2005 moisture stress was further exacerbated

in plantations due to an almost complete lack of rainfall which caused further wilting and

high rates of premature leaf loss

In the northern plantation group in August 2004 most plantations appeared to be in a good

187

state of health Trees appeared to have denser canopies than equivalent aged plantations

in the southern plantation group which suggested that the northern plantation group was

healthier In November 2004 most plantations in the northern plantation group had

received some summer rainfall which caused greater rates of canopy growth Due to

moderate rainfall the overall health in these plantations did not appear to have changed

greatly by February 2005 However by May 2005 most plantations were showing signs of

stress with increased wilting and leaf loss At no time during the study period did the

northern plantation group appear to be as moisture stressed as the southern plantation

group

Despite the apparent better health of the northern plantation group one of the more

distinct patterns to emerge from the study was that the abundances of most damage

categories were greater in the northern plantation group than the southern plantation

group Many studies show that pests and pathogens proliferate on hosts which are

stressed (Chapter 1) and it was therefore expected that the southern plantation group

would be favoured by pests and pathogens However studies also show that pests and

pathogens proliferate in more tropical environments (Beaumont 1947 Howe 1955 Krause

and Massie 1975 Stork 1988 Walklate et al 1989 Hill 1994 Nair 2001) It appears that

the higher temperatures and rainfall in the northern plantation group created conditions not

only suitable for pests and pathogens but also for greater rates of recovery in plantation

trees


Three categories of damage were identified that were accelerated by moisture stress

These included Foliar Yellowing Foliar Reddening and Physiological Necrosis The foliage

of stressed trees was observed to become discoloured either by Foliar Yellowing or Foliar

Reddening during the initial stages of senescence This damage tended to spread from the

188

tips and margins of leaves to the petiole Physiological Necrosis often occurred after Foliar

Yellowing and Foliar Reddening Premature leaf loss was also common when

Physiological Necrosis was high

Based on weather station data and other field observations February 2005 and May 2005

were identified as the two driest seasons during the survey In the southern plantation

group Physiological Necrosis was highest in February 2005 followed by May 2005 This

was expected given that these seasons were the driest Lower levels of damage in May

2005 may have been attributed to greater rates of premature leaf loss which occurred in

severely stressed trees The northern plantation group was less moisture stressed

compared with the southern plantation group and Physiological Necrosis was lower in the

northern plantation group

Levels of Foliar Yellowing and Foliar Reddening were similar in that both occurred at their

highest levels in the northern plantation group in August 2004 As expected this indicated

that these forms of damage were driven by similar climatic influences Lower levels of

damage occurred later in the survey and may have been due to greater rates of

Physiological Necrosis and premature leaf loss Effectively the yellowing and reddening

stages of leaf senescence may have led to Physiological Necrosis

Recovery from defoliation appeared to be much greater during spring and summer

compared with the colder winter months This is likely to be due to greater energy reserves

within trees during the warmer months when growth is generally greater (Bamber and

Humphreys 1965)

Interestingly no damage categories displayed clear seasonal patterns of abundance when

viewed individually however when damage was examined collectively (MDS analysis)

clear seasonal patterns were evident February 2005 and May 2005 were identified as

seasons in which collective measures of damage were the greatest

189

Effects of Drought on Pests

Despite the overshadowing effects of drought during the study drought effects also

allowed interesting insights into the effects of moisture stress on plantation trees and their

associated pests Many studies show that host stress can benefit pests by reducing host

defences (Krauss 1969 White 1984 Waring and Cobb 1992 Zangerl et al 1997

Koricheva and Larsson 1998) Phoracantha species have evolved mechanisms of

detecting stressed hosts which enables selection of lsquoweakerrsquo individuals for egg laying

(Hanks et al 1999 Lawson et al 2002) Historical moisture stress in plantation trees may

lead to greater susceptibility in the future A study by Thomson et al (2001) showed that

when E globulus was subjected to frost damage trees responded by producing new

foliage which was smaller and thinner This foliage was more susceptible to insect pests

and the phenomenon was coined lsquopost frost damage syndromersquo Similar post damage

effects have been described by Landsberg (1990a b amp c) in eucalypts suffering from

dieback

High levels of Total Insect Damage and Chrysomelid Defoliation coincided with periods of

high moisture stress in plantations in February 2005 and May 2005 This suggests that

these pests may benefit from drought conditions Many insect species prefer to feed on

soft juvenile leaves rather than tough mature leaves (Heather 1967 Tanton and Khan

1978) and the increased rate of leaf loss and regeneration may have benefited insects

through increased availability of palatable foliage

Effects of Drought on Pathogens

Many foliar pathogens require significantly humid conditions before they can infect and

sporulate on a host (Beaumont 1947 Krauss 1969) Rain also aids in the dissemination of

spores by splash dispersal (Walklate et al 1989) Before commencing the current study a

large diversity of foliar fungi were observed in plantations in southern Queensland

190

However the diversity of species appeared to decrease as the study progressed This

change may have been attributed to adversely dry conditions in southern Queensland

during drought Some fungi such as endophytic species may have benefited from the

drought because these species tend to exploit stressed hosts However few endophytes

were collected during the study period

Economic Impacts

The economic impacts of pests and pathogens in southern Queensland can only be

tentatively estimated at such an early stage in the development of the industry However a

study by Angel et al (2003) showed that the growth rate of E dunnii may be negatively

affected by pests and pathogens if the percentage of damage to the canopy exceeds

375 Elek (1997) similarly showed a threshold of 40 beyond which growth may be

compromised in other eucalypt species Given that Total Insect Defoliation reached a

maximum of 519 on one occasion and often reached 30-40 this indicates that

economic loss potentially occurred

191

5 Pests and Pathogens of Eucalypts and Hybrids A Growth Performance Trial in Southern Queensland

Introduction

The genus Eucalyptus contains over 800 species which vary greatly in form (Jacobs 1955

Brooker and Kleinig 1990) Despite the diversity of eucalypt species potentially available to

plantation growers only a handful of species have been selected for growth in plantations

(Nikles et al 2000) This is mainly because few species are suited to produce high quality

wood and have a rapid growth rate (Hollis and Brown 1987)

Eucalypts have only recently been grown in plantations in southern Queensland and there

is potential for considerable improvement within the industry In other parts of Australia

high productivity in plantations has been achieved by selective breeding of species which

are fast growing (Adams and Atkinson 1991 Eldridge et al 1994 Florence 1996 Barbour

1997 Noble 1989) A large emphasis has also been placed on resistance to pest and

pathogens (Dungey et al 1987 Lundquist and Purnell 1987 Carnegie et al 1994 Crous

and Wingfield 1996)

Species which have been successfully grown in plantations both in Australia and overseas

include E globulus E nitens E dunnii E grandis E pilularis E urophylla E maculata

E tereticornis E delegatensis E viminalis E camaldulensis E cloeziana Corymbia

maculata C citriodora and many hybrids (Lanfranco and Dungey 2001 Carnegie 2007)

Two of the more widely planted species in southern Queensland are E dunnii and E

grandis These species have become popular mainly because they are fast growing and

because there is a growing market for their wood which is used in the paper industry

Problems have emerged during the short time in which E dunnii and E grandis have been

widely planted E grandis is susceptible to both frost damage and attack by insect borers

(Phoracantha sp and Endoxyla cinerea) (Nixon and Hagedorn 1984 Manion and Zhang

192

1989 Wang et al 1998 Lawson et al 2002) E dunnii is susceptible to moisture stress

which may cause premature leaf loss (Chapter 1 amp Chapter 4 Drought in southern

Queensland)

Due to the suboptimal performance of E dunnii and E grandis plantation growers have

began to examine the performance of other eucalypt species These include E globulus

E tereticornis E camaldulensis E urophylla and their hybrids E globulus is currently the

most widely planted eucalypt species in Australia (Eldridge et al 1994 Bailey and

Duncanson 1998) E tereticornis is a fast growing species and has the largest distribution

of any eucalypt extending along the east Australian coast from southern Victoria to

northern Queensland and also New Guinea (Eldridge et al 1994) E camaldulensis occurs

in many areas of mainland Australia where it often grows along water courses

(Chippendale 1988) E camaldulensis is mainly favoured for plantations occurring in drier

areas because it has a greater drought tolerance than many species (Lanfranco and

Dungey 2001 Vinaya Rai et al 1995 Farrell et al 1996) E urophylla is native to

Indonesia and is one of only two species which is not native to Australia (the other being

E alba) E urophylla is a preferred plantation species in subtropical climates (Jǿker 2004)

Eucalypts are variable in form and many species will readily hybridise For example E

regnans (Mountain ash) and E obliqua (messmate) are co-occurring species in temperate

forests in Victoria Hybrids of these species have morphological characteristics which may

resemble either parent species or a mixture of both (Eldridge et al 1994) Such hybrids

may vary in their tolerance to climatic extremes and their susceptibility to pests and

pathogens Several natural hybrid zones in eucalypt forests have been shown to have a

greater diversity of insect and fungal species (Morrow et al 1994 Whitham et al 1994)

These areas are often called pest or pathogen lsquosinksrsquo and Whitham (1989) proposed that

they occur because hybrids are often less adapted to their environment compared with

true breeding taxa Hybrids are also more likely to suffer from stress which leads to

193

greater pest and pathogen susceptibility This is sometimes called lsquohybrid breakdownrsquo

These effects have been observed in artificial hybrids of eucalypt taxa grown in trials

alongside their parent taxa (Dungey et al 2000) Hybrids also have advantages over true

breeding taxa especially when the parent taxa are selected Fast growing species can be

crossed with species with better wood quality and greater tolerance to drought or pests

and diseases Artificial hybridisation thereby allows a degree of lsquodesignrsquo when producing

eucalypt taxa which are more suited to particular site conditions such as in plantations

(Dungey et al 2000)

Chapter Aim

In the current study a range of eucalypt species and hybrids were grown in a growth

performance trial to examine their susceptibility to pests and pathogens Seasonal

influence on pests and pathogen susceptibility was also examined


Site and Species Selection

The growth performance trial was established approximately 15 km south of Boonah in

south-east Queensland The trial was established in 1999 and the study commenced in

August 2004 when the trees were 5 years old The impacts of drought had affected the

trial by causing most tree species to prematurely drop their foliage and produce large

amounts of regrowth Tree canopies of most species therefore consisted mostly of soft

juvenile foliage rather than mature foliage

The site was relatively flat and the soil consisted of a dark brown alluvial loam which

appeared to be 1-2 m deep (roadside cutting inspection) The trial was arranged in a

randomised block design and included eight eucalypt taxa These were E dunnii E

grandis E globulus E tereticornis and the following hybrids E grandis x camaldulensis

194

E tereticornis x urophylla E urophylla x camaldulensis and E urophylla x grandis (Table

51) All of these species were grown from seed which was collected from parent stock (no

clones were used) Each taxon was grown in three separate blocks consisting of 6 rows of

12 trees (72 trees per block) The spacing of the trees was 2 m between stems within rows

and 4 m between rows The area of each block was approximately 0057 ha All blocks

were arranged randomly and surrounded on all sides by an equal aged mixed-species

plantation of E dunnii and E grandis (Figure 5-1)

Figure 5-1 A representation of the taxa trial layout (marked with a square) Different coloured dots within the square represent trees belonging to different taxa The blocks of taxa were grown side by side and arranged randomly (not to scale) The trial was surrounded on all sides by even aged E dunnii plantation trees

195

Species Native Range Morphology and Ecology

E dunnii

(Dunnrsquos white gum)

Two relatively small populations occur in northern NSW which are 120 km apart (Boland et al 1984 Benson and Hager 1993 Specht et al 1995) Because these populations are estimated to occupy less than 80000 ha the species is listed as endangered (Briggs and Leigh 1988)

Tree to 50 m Bark grey to grey-brown fibrous-flaky on lower trunk smooth above white or grey shedding in short ribbons Juvenile leaves opposite orbiculate to ovate cordate dull grey-green Adult leaves disjunct narrow-lanceolate or lanceolate wide green dull concolorous Buds ovoid Fruit hemispherical or conical or campanulate (Brooker and Kleinig 1999)

Prefers fertile basaltic and alluvial soils on the margins of rainforests (Booth and Jones 1988 Booth et al 1999 Jovanovic et al 2000)

E grandis

(Flooded gum)

Numerous populations occur on the east Australian coast from Newcastle (northern NSW) to Bundaberg (southern QLD) (Angel 1999 Jovanovic et al 2000 Wang et al 1998)

Tree to 50 m in height Bark rough at the base fibrous or flaky grey to grey-brown Leaves stalked lanceolate to broad lanceolate discolorous Flowers White

Prefers deep alluvial and volcanic loams with high moisture such as in valleys and flats

E globulus

(Blue gum)

Extensive populations occur in Tasmania the Bass Strait Islands and south-eastern Victoria (Eldridge et al 1994)

Tree to 45 m Bark smooth apart from base which has persistent slabs shedding in large strips and slabs smooth bark white cream grey yellowish or pale creamy orange often with ribbons of decorticated bark in the upper branches Juvenile stem square in cross-section and winged Juvenile leaves opposite and sessile for many pairs oblong to elliptical then ovate or broadly lanceolate upper surface green or slightly glaucous and the lower surface copiously white-waxy Adult leaves alternate lanceolate to falcate (Brooker and Kleinig 1999)

Prefers a range of soil conditions from gradational clay loams to uniform sands with mean annual rainfall ranging from 650 to 1000 mm (Weston et al 1991)

E tereticornis

(Forest red gum)

E tereticornis has the largest distribution of any eucalypt species which extends along the east Australian coast from south-east Victoria through New South Wales and Queensland and also occurs in New Guinea (Brooker and Kleinig 1999)

Tree to 50 m usually much smaller in exposed coastal situations (Alverado et al

2006) Bark smooth white grey shedding in large flakes Adult leaves disjunct narrow ovate and falcate glossy green 10-20 cm long and 1-3 cm wide Flowers white and in some areas pink appearing June to November Fruit ovoid with raised disc

Prefers hind dunes along coastal streams and wet sclerophyll forests (Brooker and Kleinig 1999)

Table 51 Species characteristics of the eucalypt taxa (some of which were hybridised)

196


E camaldulensis

(River red gum)

Populations occur in most areas of mainland Australia except southern Western Australia south-western South Australia and the eastern coastal areas of Queensland New South Wales and Victoria (Chippendale 1988)

Tree to 30 m (Bren and Gibbs 1986) although some authors (eg Boland 1984 Brooker and Kleinig 1999) record trees to 45 m Bark smooth mottled and periodically shedding through the year while becoming rough at the base Leaves petiolate to broadly lanceolate Hemispherical buds on stalks (Brooker and Kleinig 1999)

Prefers the edges of rivers where its roots have access to water (Brooker and Slee 1996)

E urophylla

(Timor mountain gum)

E urophylla is native to south east Indonesia where it occurs on the islands of Timor Flores Wetar Lembata (Lomblem) Alor Adonara and Pantar The two main centres are Timor and Flores (Jǿker 2004)

Tree to 45 m tall Bark variable depending on moisture and altitude usually persistent and subfibrous smooth to shallow close longitudinal fissures red-brown to brown sometimes rough especially at the base of the trunk Juvenile leaves subopposite stalked broadly lanceolate adult leaves phyllodinous subopposite to alternate long stalked broadly lanceolate discolourous lateral veins just visible Buds semi-circular black Flowers peduncle somewhat flattened 8-22 mm long (Jǿker 2004)

Prefers wet soils with loose texture soil (volcanically derived) and occurs in dry and wet tropical forest (Jǿker 2004)

Hybrid taxa

E grandis x camaldulensis

E tereticornis x urophylla

E urophylla x camaldulensis and

E urophylla x grandis

None of the parent taxa of the hybrids are known to hybridise under natural conditions and therefore no native geographical ranges occur

Many hybrids have phenotypic characteristics which are a blend of the parent taxa However the resemblance of the hybrids to either parent taxa may vary greatly between individuals (Eldridge et al 1994)

Identifying and Measuring Damage

Damage was assessed using a modified version of the lsquoCrown Damage Index

Assessmentrsquo by Stone et al (2003) also described in Chapter 4 (Table 41) Each taxon

was assessed by examining levels of damage on the inner six trees of each block Three

blocks were sampled for each species so that 18 individual trees were assessed for each

taxon during each round of sampling (Figure 5-2) All damage less than 10 was referred

197

to as low damage between 10 and 20 was referred to as moderate and damage

above 20 was referred to as high

Sampling Regime

The trial was assessed in August 2004 November 2004 February 2005 and May 2005

Climate Data

Rainfall data were obtained from the Australian Bureau of Meteorology for the Amberley

weather station which occurred approximately 15 km from the site (Chapter 4 Figure 4-5

and Figure 4-6)

Multivariate Analyses

Multivariate analyses were carried out using the Primer 5 statistical package (Clarke and

Gorley 2001) The Bray-Curtis similarity coefficient was employed to construct a similarity

matrix from the log (n+1) transformed values ( Total Damage for each damage category

within each taxa between seasons) This matrix was then subjected to non-metric

Six trees selected within each block for the assessment

Figure 5-2 A graphical representation of the six trees (red dots) selected for damage assessment within each block of the trial

198

multidimensional scaling (MDS) ordination One way crossed Analysis of Similarities

(ANOSIM) (Clarke and Gorley 2001) were carried out to ascertain whether the

compositions of the damage categories differed significantly between taxa and between

seasons The factors employed in each of the tests are specified in the results In each

test the null hypothesis that there were no significant differences among groups was

rejected if the significance level (p) was lt5 The R statistic value was used to ascertain

the extent of any significant differences (Clarke and Gorley 2001) Any R values lt01 were

regarded as negligible Where ANOSIM detected a significant difference among priori

groups and the R-statistic was gt01 similarity percentages (SIMPER) (Clarke and Gorley

2001) were used to identify which damage categories made the greatest contribution to

those differences

Results

Average Measures of Damage

Eleven damage categories were identified and examined (Table 52) Most of the damage

recorded during the survey was caused by insects Total Insect Defoliation caused the

highest Total Damage Given that most of the damage within this category was caused

by chrysomelid beetles it is not surprising that the second highest Total Damage was

caused by Chrysomelid Damage The highest measures of damage after these were Total

Insect Necrosis Total Fungal Damage Foliar Yellowing and Physiological Necrosis All

other damage occurred at relatively low levels (Table 52)

199

Damage category of Damage Rank

Total Defoliation 3042 1st








Teratosphaeria Damage 017 9th


Mirid Damage 0001 11th

Total 100

Comparing Taxa

Multivariate statistics were used to show how measures of damage varied collectively (all

damage categories) between the different taxa Pest and disease species were therefore

compared both as assemblages and as individual categories A one way crossed analysis

of similarities showed that collective levels of damage varied significantly between some

but not all taxa (significant when Plt005) (Table 53) No significant differences were

observed for E tereticornis E tereticornis x urophylla or E urophylla x camaldulensis E

dunnii was significantly different from E globulus E grandis x camaldulensis E

tereticornis and E urophylla x camaldulensis while E grandis was significantly different

from E grandis x camaldulensis and E urophylla x camaldulensis (Table 53) Significant

R values (gt01) which ascertain the extent of differences between collective measures

indicated that E grandis x camaldulensis followed by E globulus and E dunnii were the

most different species in the trial in terms of collective measures of damage Because the

Global R value of the analysis (0408) was less than 05 this infers that that there is a

generally significant difference between all taxa in terms of collective measures of

damage


Table 52 Total Damage and rank (1st-11

th) caused by

each damage category for all measurements (all taxa)

200

damage categories made the greatest contribution to differences between taxa in terms of

collective measures of damage (Table 54) These were Total Defoliation Chrysomelid

Defoliation Total Insect Necrosis Total Fungal Damage Foliar Yellowing Phylacteophaga

Blisters and Foliar Wasp Galls

201

All Species (P=01 Global R=0408)

E dunnii E globulus E grandis E tereticornis E grandis x camaldulensis


E urophylla x camaldulensis

P R P R P R P R P R P R P R E dunnii E globulus 01 0575 E grandis 27 0235 01 0492 E tereticornis 04 032 01 0679 48 0278 E grandis x camaldulensis 01 065 03 0394 01 0352 05 0519 E tereticornis x urophylla 40 0191 01 0648 33 0287 675 -0056 01 0648 E urophylla x camaldulensis 03 0383 01 0796 05 05 155 0148 01 0824 595 -0037 E urophylla x grandis 12 0298 01 0633 45 025 286 0065 04 0472 200 0102 127 0157

Pure Taxa

Rank E dunnii E globulus E grandis E tereticornis

1st

2nd

3

rd

4th

5th

Total Defoliation (290) Chrysomelid Defoliation (289) Total Insect Necrosis (00) Total Fungal Damage (62) Foliar yellowing (52)

Total Defoliation (152) Chrysomelid Defoliation (152) Total Insect Necrosis (203) Total Fungal Damage (116) Phylacteophaga Blisters (44)

Chrysomelid Defoliation (122) Total Defoliation (122) Total Insect Necrosis (93) Total Fungal Damage (49) Foliar yellowing (42)


Hybrid Taxa

Rank E grandis x camaldulensis E tereticornis x urophylla E urophylla x camaldulensis E urophylla x grandis

1st

2nd

3

rd

4th

5th

6th

Total Defoliation (117) Chrysomelid Defoliation (117) Total Insect Necrosis (114) Total Fungal Damage (21) Phylacteophaga Blisters (00) Foliar wasp galls (00)

Total Defoliation (271) Chrysomelid Defoliation (271) Total Insect Necrosis (96) Total Fungal Damage (82)



Table 53 Significance levels (p) and R ndashstatistic values for both global and pair-wise comparisons in a one way ANOSIM test of all damage

categories in all 8 taxa Significant results in bold (Plt01 Rgt05) (values in bold with asterix are significant)

Table 54 Damage categories detected by SIMPER as those most responsible for typifying the damage for each of the Eucalyptus species and hybrids Samples collected in the different seasons have been pooled in this analysis Mean percentage () of damage included in brackets

202

Multi Dimensional Scaling (MDS) using ordination (dissimilarity by distance) was also used

to examine and compare collective measures of damage between taxa (Figure 5-3) This

analysis showed indistinct separation by distance of most taxa E globulus showed some

isolation by distance in the analysis but clustered into two distinct groups (G1 and G2)

This indicated that differences in terms of collective measures of damage occurred

between these two groups Similar separation by distance with double groupings was also

observed for E grandis x camaldulensis (G3 and G4)

The stress value (lt2) indicated that the ordination was an acceptable representation of the

observed variability between the measurements in the analysis The ordination supported

what was suggested by ANOSIM namely that E globulus and E grandis x camaldulensis

were the most different taxa within the trial in terms of collective measures of damage

Effects of Seasonal Climate

The taxa trial occurred within 20 km of the southern plantation estate as discussed in

Figure 5-3 Two dimensional MDS ordination of the second-stage similarity matrix containing the correlations between each pair of similarity matrices constructed from the severity of all 11 damage category all samples (8 species rated in 4 seasons) The points are coded for eucalypt species

G1 G2

G3

G4

203

Chapter 4 Seasonal trends in regional climate between these two areas were very similar

The weather data presented in Chapter 4 (Figure 4-5 and Figure 4-6) was used to make

inferences about variability in levels of damage between seasons


damage categories) between different seasons of sampling across all taxa (pest and

disease species being compared as assemblages) A one way crossed analysis of

similarities (ANOSIM) showed that collective levels of damage varied significantly (Plt05)

between all four seasons of sampling (Table 55) R values from this analysis indicated

that the most different season in terms of collective measures of damage was August

2004



P R P R P R P R

Aug 04

Nov 04 01 0763

Feb 05 01 0634 01 0271

May 05 01 0757 01 0481 01 0562



terms of collective measures of damage Total Defoliation Chrysomelid Defoliation Total

Fungal Damage and Total Insect Necrosis were ranked among the highest contributors

(Table 56)

Rank

Seasons of Sampling


1st

2nd

3

rd

4th

5th

Total Defoliation (201) Chrysomelid Defoliation (201) Total Fungal Damage (166) Total Insect Necrosis (134) Foliar Yellowing (72)

Total Defoliation (161) Chrysomelid Defoliation (161) Total Fungal Damage (37) Teratosphaeria Damage (05)


Total Defoliation (300) Chrysomelid Defoliation (300) Total Insect Necrosis (101) Total Fungal Damage (46) Foliar Yellowing (35)

Table 55 Significance levels (p) and R ndashstatistic values for both global and pair wise comparisons in a one way ANOSIM test of all damage categories in the four seasons of sampling Significant results (Rgt01)

Table 56 Damage categories detected by SIMPER as those most responsible for typifying the damage for each season Mean percentage () of damage included in brackets

204


to examine and compare collective measures of damage between seasons (Figure 5-4)

The MDS showed a distinct separation by distance of the points representing collective

measures of damage for August 2004 and very little separation for November 2004

February 2005 and May 2005 which grouped together The stress value (lt2) indicated that

the ordination was an acceptable representation of the observed variability between the

measurements in the analysis The ordination supported what was suggested by ANOSIM

that August 2004 was the most different season followed by November 2004 February

2005 and May 2005 Greater separation was observed for this ordination than from the

previous analysis comparing different taxa (Figure 5-4)

Total Defoliation and Chrysomelid Defoliation

The majority of defoliation was caused by chrysomelid beetles and therefore patterns of

abundance for Total Defoliation (Figure 5-5) and Chrysomelid Defoliation (Figure 5-6) were

very similar The abundance of damage by these damage categories varied greatly

Figure 5-4 Two dimensional MDS ordination of the second-stage similarity matrix containing the correlations between each pair of similarity matrices constructed from the severity of all 10 damage category all samples (8 species rated in 4 seasons) The points are coded for season The analysis shows two groupings

205

between individual trees individual taxa and also between seasons

Relatively low levels of Chrysomelid Damage occurred on E dunnii compared with other

taxa at the beginning of the survey however levels of damage increased successively

with each season thereafter E globulus and E grandis exhibited moderate levels of

damage at the beginning of the survey which fluctuated slightly with each season E

grandis x camaldulensis showed the opposite pattern of abundance by exhibiting

decreasing levels of damage as the survey progressed E tereticornis E tereticornis x

urophylla and E urophylla x camaldulensis showed relatively high levels of damage during

the first half of the survey and then much higher levels of damage during the second half

Similarly E urophylla x grandis had moderate levels of damage during the first half of the

survey and then much higher levels during the second half

206

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

Figure 5-6 Mean percentages of Chrysomelid Defoliation (plusmn SE) for each eucalypt taxon in four seasons of sampling (A) August 2004 (B) November 2004 (C) February 2005 and (D) May 2005)


A B C D

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

Figure 5-5 Mean percentages of Total Defoliation (plusmn SE) for each eucalypt taxon in four seasons of sampling (A) August 2004 (B) November 2004 (C) February 2005 and (D) May 2005)


A B C D

207


Total Damage for Total Insect Necrosis was moderately high (lt20) for most taxa

during most seasons with the exception of high levels occurring on E globulus in

November 2004 and February 2005 (Figure 5-7) High levels were also observed on E

urophylla x camaldulensis in November 2004 and May 2005

Total Fungal Damage

In August 2004 most taxa were affected by Total Fungal Damage (Figure 5-8) However

by November 2004 levels of damage had decreased dramatically In February 2005

levels increased again on E tereticornis E tereticornis x urophylla E urophylla x

camaldulensis and E urophylla x grandis before decreasing again in May 2005 Damage

remained low on E dunnii E globulus and E grandis in February 2005 before increasing

again in May 2005 Levels of damage were consistently low on E grandis x camaldulensis

during all seasons of sampling

208

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

Figure 5-7 Mean percentages of Total Insect Necrosis (plusmn SE) for each eucalypt taxon in four seasons of sampling (A) August 2004 (B) November 2004 (C) February 2005 and (D) May 2005)


A B C D

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x granEucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

Figure 5-8 Mean percentages of Total Fungal Damage (plusmn SE) for each eucalypt taxon in four seasons of sampling (A) August 2004 (B) November 2004 (C) February 2005 and (D) May 2005)


A B C D

209

Foliar Yellowing

In August 2004 all taxa with the exception of E globulus and E tereticornis x urophylla

were affected by low to moderate levels of Foliar Yellowing (Figure 5-9) Damage was

completely absent from all taxa in November 2004 E dunnii E grandis E urophylla x

camaldulensis and E urophylla x grandis were affected by low levels of damage in

February 2005 E tereticornis and E urophylla x grandis exhibited low levels of damage in

May 2005 while E tereticornis x urophylla exhibited high levels of damage


Physiological Necrosis was completely absent in all taxa during the survey until the final

seasonal sample in May 2005 when E dunnii E globulus and E grandis were affected by

high levels of damage and E grandis x camaldulensis E tereticornis x urophylla and E

urophylla x camaldulensis were affected by low levels of damage (Figure 5-10)

210

Figure 5-9 Mean percentages of Foliar Yellowing (plusmn SE) for each eucalypt taxon in four seasons of sampling (A) August 2004 (B) November 2004 (C) February 2005 and (D) May 2005)


00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt TaxaP

erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)A B C D

Figure 5-10 Mean percentages of Physiological Necrosis (plusmn SE) for each eucalypt taxon in four seasons of sampling (A) August 2004 (B) November 2004 (C) February 2005 and (D) May 2005)


00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

211

Remaining Damage Categories

All remaining damage categories including Phylacteophaga Blisters Foliar Wasp Galls

Mirid Damage Teratosphaeria Damage and Scale Insect Damage caused negligible

damage throughout the study period (Table 56 Table 57 Table 58 and Table 59)

212

Aug-04

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 92 196 97 326 264 313 292 149

plusmn SE 07 14 17 34 00 23 47 21


M 92 196 94 326 250 313 292 149

plusmn SE 60 60 42 42 42 42 42 42


M 132 120 88 163 183 219 198 167

plusmn SE 17 13 19 24 23 18 21 23


M 09 172 14 00 00 00 00 00

plusmn SE 21 201 24 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Mycosphaerella Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Fungal Damage

M 144 150 94 288 83 167 410 128

plusmn SE 17 11 28 45 29 15 37 19

Foliar Yellowing

M 174 00 139 56 42 00 56 111

plusmn SE 63 00 77 56 42 00 56 77

Scale Insect Damage

M 00 00 03 00 00 00 00 00

plusmn SE 00 00 05 00 00 00 00 00

Table 56 Mean (M) Standard error (SE) and proportion of total damage () for each lsquodamage categoryrsquo and each eucalypt taxon during August 2004

213

Nov-04

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 264 135 205 178 104 188 354 128

plusmn SE 19 12 21 29 07 09 51 08


M 264 135 25 177 14 188 354 128

plusmn SE 60 60 42 42 42 42 42 42


M 214 384 125 104 83 125 208 87

plusmn SE 15 25 00 14 14 00 51 07


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 51 00 00 00 00 00 00

plusmn SE 00 100 00 00 00 00 00 00

Total Fungal Damage

M 00 121 42 21 00 00 111 00

plusmn SE 00 34 23 07 00 00 51 00

Foliar Yellowing

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Table 57 Mean (M) Standard error (SE) and proportion of total damage () for each lsquodamage categoryrsquo and each eucalypt taxon during November 2004

214

Feb-05

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 373 167 146 417 125 458 436 413

plusmn SE 22 12 07 34 05 24 25 30


M 373 161 146 417 125 458 431 413

plusmn SE 60 60 42 42 42 42 42 42


M 175 239 146 153 125 167 156 125

plusmn SE 08 20 07 13 00 15 17 00


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 00 03 00 00 00 00 00

plusmn SE 00 00 06 00 00 00 00 00

Total Fungal Damage

M 30 00 10 340 00 292 188 264

plusmn SE 08 00 06 43 00 15 37 47

Foliar Yellowing

M 36 00 28 00 00 00 14 56

plusmn SE 29 00 19 00 00 00 14 33

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Table 58 Mean (M) Standard error (SE) and proportion of total damage () for each lsquodamage categoryrsquo and each eucalypt taxon during February 2005

215

May-05

E dunnii

E globulus

E grandis

E tereticornis






M 307 416 340 00 07 63 10 00

plusmn SE 66 65 85 00 05 16 11 00

Total Defoliation

M 429 117 219 244 66 417 549 444

plusmn SE 22 12 37 21 04 15 27 15


M 429 116 219 243 66 417 549 444

plusmn SE 60 60 42 42 42 42 42 42


M 98 70 101 69 63 63 267 163

plusmn SE 08 03 15 05 00 00 38 27


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 111

plusmn SE 00 00 00 00 00 00 00 192


M 02 00 00 00 00 00 00 00

plusmn SE 04 00 00 00 00 00 00 00

Total Fungal Damage

M 73 193 49 03 00 00 10 10

plusmn SE 21 23 27 04 00 00 06 06

Foliar Yellowing

M 00 00 00 14 00 333 83 42

plusmn SE 00 00 00 14 00 123 58 23

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Table 59 Mean (M) Standard error (SE) and proportion of total damage () for each damage category and each eucalypt taxon during May 2005

216

Discussion


Due to the close proximity of the taxa trial to the southern plantation group discussed

in Chapter 4 the same weather data (BOM) were used in this study to examine the

influence of season on pests and pathogens Like other plantations within the region

the taxa trial was subjected to severe drought conditions (2001-2006) which caused

trees to become moisture stressed As a result the foliage of many taxa was

observed to suffer premature leaf loss (abscission) during especially dry periods

After rain these trees often responded quickly by producing copious amounts of new

foliage (flush growth) These effects may have overshadowed the effects of pests

and pathogens in the trial and made it very difficult to attribute changes in damage to

actual changes in the size of insect and fungal populations

Two damage categories were identified as being under seasonal influence These

were Total Fungal Damage and Physiological Necrosis All other damage categories

showed erratic variability in damage levels both between taxa and season Total

Fungal Damage was most severe on the majority of taxa in August 2004 and

February 2005 These months coincided with similarly low rainfall Studies show that

although pathogens tend to proliferate during humid conditions (Beaumont 1947

Krausse 1975 Daniel and Shen 1991 Agrios 2005) they may also benefit from dry

conditions if it causes their host to become stressed (reducing defences) (Bertrand et

al 1967 Yarwood 1959 Colhoun 1973 Hepting 1963 Boyer 1995 Schoenweiss

1975 1981) Observations of the general health of the taxa trial in August 2004 and

February 2005 indicated that the trees were stressed which may have led to a

greater proliferation of foliar pathogens and hence greater levels of Total Fungal

217

Damage however this conflicts with observations made in other plantations during

the survey which indicated that many pathogens were negatively affected by dry

conditions Another possibility for the greater levels of Total Fungal Damage is a

dilution caused by flush growth in November 2004 and May 2005 which may have

reduced the proportion of damaged leaves in canopies A general trend of

decreasing Total Fungal Damage was observed during the survey which may have

been due to increasingly adverse dry conditions

Observations in the field indicated that Physiological Necrosis occurred when trees

became stressed This is consistent with the appearance of damage on many taxa in

May 2005 because very low rainfall occurred from January to April 2005 These

effects are also consistent with other studies (Old 1990 Vinaya Rai et al 1995

McGrath 1999)

Foliar Yellowing was observed on most taxa Foliar Yellowing had multiple causes

such as insect and fungal damage or the early development of Physiological

Necrosis Yellowing can also be caused by nutrient deficiencies (Graham and Webb

1991 Dell and Malajczuk 1994) Given that under the right conditions yellowing

could arise very quickly it is difficult to definitively attribute the observed variability in

yellowing to seasonal affects

August 2004 was identified as being the most different season in terms of collective

measures of damage This may have been due to greater levels of Total Fungal

Damage Foliar Yellowing Phylacteophaga Blisters and Scale Insect Damage on

most taxa during this time Given that August was a period of extremely low rainfall

in southern Queensland it was expected that this season would have a strong

influence on pests and pathogens Greater levels of yellowing were expected due to

the likelihood of greater moisture stress in plantations Higher levels of Total Fungal

218

Damage were unexpected because it was thought that this would occur during

summer when high temperatures and high rainfall lead to high humidity

Comparing Taxa

The majority of Total Defoliation was caused by chrysomelid beetles which were

abundant during most stages of the survey Many chrysomelid species prefer soft

juvenile foliage to adult foliage (de Little and Madden 1975 Tanton and Khan 1978)

The large amounts of flush growth produced by taxa during periods of high moisture

stress may have benefited the development of feeding chrysomelid beetles Levels

of damage appeared to generally increase between seasons which may have been

due to growth of the chrysomelid population over time It is interesting to note that

although damage levels increased on most species damage on E globulus E

grandis and E camaldulensis x grandis remained relatively low This may indicate

that these species have greater resistance to chrysomelid attack or a greater rate of

recovery Observations in the field suggested these species were less affected by

premature leaf loss during dry conditions which may have led to less epicormic

growth and less feeding by chrysomelids

Total Insect Necrosis was similarly high on all taxa during the survey Greater levels

of damage occurred on E globulus in November and observations in the field

indicated that most of this damage was caused by a single sap-sucking insect

species Platybrachys sp (Eurybrachidae) This species caused small interveinal

necrotic patches on foliage during feeding and also scars on the stems from the

oviposition of eggs

Physiological Necrosis mostly occurred at low levels and was most severe on E

dunnii E globulus and E grandis Although this suggests that these species are

219

more susceptible to moisture stress this is in conflict with field observations While

examining E dunnii E globulus and E grandis it was observed that all other taxa

within the trial had prematurely dropped more foliage during periods of high moisture

stress When rating the incidence and severity of Physiological Necrosis taxa with

more foliage generally had greater levels of Physiological Necrosis Because

premature leaf loss is likely to be a better indicator of moisture stress than

Physiological Necrosis E dunnii E globulus and E grandis should be considered to

be less susceptible to moisture stress

Conclusion

The 2001-2006 drought had an impact on both the taxa trial and its associated

diseases and pests It was difficult to make inferences regarding the susceptibility of

taxa to diseases and pests while they were stressed This problem was exacerbated

by the effects of leaf loss and regeneration which made it very difficult to attribute

changes in measures of damage to actual changes in the size of insect and fungal

populations For example a tree with a moderate level of infection by a pathogen

may appear to be more severely affected once foliage is prematurely lost or

conversely the same tree may appear healthier after the production of flush growth

despite no actual change in the number of infected leaves It must therefore be

stated that these effects had the potential to affect all measures of damage and

undermine the interpretation of the findings of the study

Despite the overshadowing effects of drought some patterns were observed which

allowed inferences to be made regarding the susceptibility of taxa to moisture stress

Given that no taxa showed consistency in their susceptibility to pests and diseases

between seasons this suggested that susceptibility may be under greater influence

of external factors such as climate Fluctuations in the abundance of pests and

220

pathogens were erratic and this indicated that lsquoshort term effectsrsquo such as rainfall

events may have a greater influence on host susceptibility than long term

accumulative effects or inherent susceptibility Observations in the field indicated that

trees prematurely lost their foliage very quickly during dry conditions and then

produced flush regrowth quickly after rain These processes are likely to be the main

cause of erratic variability in damage levels

It is important to note that the susceptibility of eucalypts to pests and pathogens may

vary depending on site conditions Due to influences such as lsquomonoculture effectsrsquo

the performance of eucalypt species in the taxa trial may be different to that if they

were grown in a plantation In the absence of lsquochoicersquo some pests may simply utilise

the only resource available to them (Kavanagh and Lambert 1990) Overall the trial

suggested that the most suitable tree species for growth in plantations in southern

Queensland were E dunnii E grandis and possibly even E globulus

221

6 The Pathogenicity of Fungi Associated with Stem Basal Cankers of Eucalypt Plantations

Introduction

In 2003 plantation growers in southern Queensland had increasing concerns about

the occurrence of stem basal cankers in one and two-year-old plantations The

cankers superficially consisted of dark brown swellings at the base of trees These

swellings often produced kino when heavily cracked and the removal of bark

revealed necrosis of the vascular cambium (Figure 6-1) Several fungal species were

isolated from cankers including saprophytes such as Pestalotiopsis sp Alternaria

sp and Fusarium sp Opportunistic pathogens which were isolated included

Cytospora eucalypticola Neofusicoccum ribis and Holocryphia eucalypti

Most Cytospora species are considered to be weakly pathogenic species which may

cause small superficial cankers on branches and stems of eucalypt hosts (Fraser

and Davidson 1985 Old et al 1986 1990 Fisher et al 1993 Yuan and Mohammed

1997 Old and Davison 2000 Adams et al 2005 Carnegie 2007a) Cytospora

eucalypticola is the most commonly isolated species in eucalypt plantations (Old et

al 1986 Old and Davison 2000) As well as being weakly pathogenic C

eucalypticola has both endophytic and saprophytic characteristics Bettucci et al

(1999) found that C eucalypticola was commonly isolated from healthy stems of E

grandis in the absence of a disease response Yuan and Mohammed (1997) found

C eucalypticola to be commonly associated with stressed hosts such as roadside

trees suffering from crown dieback Old et al (1991) isolated the fungus from dead

lower branches of E nitens and E globulus in plantations in Tasmania

The genus Botryosphaeria contains 16 species for which Botryosphaeria dothidea is

222

the lectotype (Cesati and De Notaris 1963 Barr 1972) B ribis was considered to be

synonymous with B dothidea until it was differentiated based on combined multiple

gene genealogies and phenotypic characters by Slippers et al (2004) A revision of

the Botryosphaeriaceae has renamed B ribis as Neofusicoccum ribis (Crous et al

2006) N ribis may cause a range of symptoms on eucalypts including dieback stem

bleeding necrosis coppice failure and cankers (Davison and Tay 1983 Smith et al

1994 Old and Davison 2000 Burgess and Wingfield 2002) The species is also an

endophyte of healthy hosts and may become pathogenic and cause disease in

stressed hosts It is therefore commonly referred to as a latent pathogen (Old et al

1990 Fisher et al 1993 Zhonghua et al 2001 Burgess et al 2004 Slippers et al

2004) Pathogenicity tests on E delegatensis showed that N ribis is more

pathogenic than C eucalypticola N ribis has also been isolated from wood

associated with the galleries of wood borers such as Cerambycidae (Fraser and

Davison 1985) Whyte (2002) found a Fusicoccum anamorph of Neofusicoccum

associated with foliar lesions of E camaldulensis which also occurs in association

with a parasitic-wasp species (Eulophidae)

Holocryphia eucalypti (Gryzenhout et al 2006) previously known as Cryphonectria

eucalypti Endothia gyrosa (Venter et al 2001 2002) and Endothia havanensis

(Davison 1982 Davison and Tay 1983 Fraser and Davison 1985) is a canker

pathogen that causes various levels of damage to at least 20 species of eucalypts in

a range of localities across Australia South Africa and Uganda (Davison 1982

Fraser and Davison 1985 Walker et al 1985 Old et al 1986 Davison and Coates

1991 White and Kile 1993 Yuan and Mohammed 1997a Wardlaw 1999

Gryzenhout et al 2003 Gryzenhout et al 2006) A recent study showed that H

eucalypti is also pathogenic to Tibouchina urvilleana which is currently the only

223

known non-eucalypt host (Heath et al 2007) H eucalypti is particularly widespread

in eastern Australia where it is a common canker pathogen of eucalypts (Walker et

al 1985 Old et al 1986 Yuan and Mohammed 1997a Wardlaw 1999 Carnegie

2007a 2007b) Although once thought to occur in North America the species was

eventually shown to be a different species based on phylogenetic analysis (Shear et

al 1917 Stipes and Phillips 1971 Appel and Stipes 1986 Roane et al 1974 Venter

et al 2001 2002) Symptoms of the disease are variable and may include bark

cracks cankers kino exudation and dieback of coppice shoots branches and stems

(Old et al 1986 Walker 1985) Reports also show that symptoms vary between

localities For example fruiting bodies of the teleomorph are commonly associated

with eucalypts in Tasmania (Yuan and Mohammed 1997a) whereas only the

Endothiella anamorph has been observed in Western Australia (Shivas 1989

Shearer 1994 Jackson et al 2004) Infections have been shown to be facilitated by

wounding of the host such as by cracks and fissures in the stem such as damage

cause by wind (Yuan 1998 Yuan and Mohammed 2001 Ferreira and Milani 2002)

Pathogenicity studies have shown that the species is a mild pathogen which is

capable of killing seedlings and stressed trees (Walker et al 1985 Old et al 1986

Yuan and Mohammed 1997 Wardlaw 1999 Gryzenhout et al 2003 Carnegie

2007a 2007b Heath et al 2007) Hosts which are stressed due to repeated

defoliation by insects may be at greater risk of infection (Old et al 1990) Gryzenhout

et al (2003) showed that different clones of E grandis vary in their susceptibility to

H eucalypti The pathogenicity of the species can also vary between isolates (Yuan

and Mohammed 1999)

When isolating fungi from cankers of diseased tree hosts it is common to isolate

more than one species This appears to be particularly common in stressed hosts

224

because opportunistic species such as saprophytes latent pathogens and primary

pathogens may be associated as assemblages (Yuan and Mohammed 1997

Bettucci et al 1999 Burgess et al 2004) When isolating fungi from basal cankers in

southern Queensland several fungal species including saprophytes latent

pathogens and opportunistic pathogens were collected (Hardy and Burgess 2003

pers comm) Very few studies have examined the interactions of different pathogens

in association with the same host (in vivo) however it has long been recognised that

some fungi can produce chemicals which reduce the growth of other species in vitro

This is commonly observed when stored fungal colonies become contaminated with

ubiquitous species such as Penicillium which can inhibit the growth of other fungal

species (Wainwright and Swan 1986) Fungal interactions are likely to vary

depending on the species involved

Three testable hypotheses describe the interactions of canker pathogens within a

living host These are

1) Antagonism whereby one pathogen reduces the pathogenicity of another

pathogen and causes a reduced disease response

2) Synergism whereby one pathogen increases the pathogenicity of another

pathogen and causes a greater disease response and

3) No effect whereby pathogens do not influence the pathogenicity of other

pathogens and the disease response is unaffected

Chapter Aim

The aim of this study was to test hypotheses 1 2 amp 3 by infecting eucalypt hosts with

three canker pathogens in various combinations and examining the resulting disease

response Cytospora eucalypticola Neofusicoccum ribis and Holocryphia eucalypti

225

were selected because they are all considered to be opportunistic pathogens which

mainly affect stressed eucalypts Based on other studies it was expected that H

eucalypti would be the most pathogenic species followed by N ribis and then C

eucalypticola (Old et al 1986 Old and Davison 2000) Pathogenicity experiments

were conducted in summer and winter to examine seasonal effects on disease

expression

Figure 6-1 A typical basal canker of a two-year-old plantation eucalypt (E dunnii) Symptoms include darkening of the bark from grey to brown at the base (stocking) severe necrosis of the vascular cambium beneath the bark and longitudinal cracking of the bark surface

Cracking of the bark

Darkening of the bark

Margin of healthy and diseased

tissue

Healthy section of vascular cambium

Diseased section of vascular cambium

226


Collection and Isolation

Opportunistic collecting of pathogens was conducted in several plantations in

southern Queensland over a two year period and diseased material was collected

from approximately 50 trees during this time Diseased material was collected by

stripping bark from diseased stems using a sterile knife to locate the disease margin

and then chipping sections of diseased wood into a paper bag using a sterile

machete Specimens were refrigerated until they could be examined later in the

laboratory (generally within 5-10 days) Wood chips were then cut into smaller pieces

under sterile conditions and surface sterilised with alcohol and flamed for two-three

seconds (Old et al 1986) The pieces were then placed onto Petri-dishes containing

half strength potato dextrose agar (PDA) and incubated in the dark at 25C for three

to four days The resulting fungal cultures were then subcultured onto fresh PDA

plates and maintained at 25 C Fresh subcultures were made every few months to

keep cultures uncontaminated and in a state of active growth Long-term storage of

cultures was achieved by placing a 1 cm cube of myceliaagar in a sterile sealed vial

of distilled water which was then stored at 15 C

Species Identification

Molecular and classical taxonomy were used to identify fungi When identifying

specimens using molecular techniques the culture was first grown on 2 (wv) PDA

at 20C in the dark for 4 weeks Mycelium from the culture was then harvested using

a sterile razor blade and placed in a 15 ml sterile Eppendorfreg tube The mycelium

was then frozen in liquid nitrogen ground to a fine powder and genomic DNA was

227

extracted A part of the internal transcribed spacer (ITS) region of the ribosomal DNA

operon was amplified using the primers ITS-1F (5rsquo CTT GGT CAT TTA GAG GAA

GTA A) Gardes and Bruns (1993) and ITS-4 (5rsquoTCC TCC GCT TAT TGA TAT GC 3rsquo)

(White et al 1990)

To compare DNA sequences of fungal species with other closely related species

additional ITS sequences were obtained from GenBank Sequence data were

assembled using Sequence Navigator version 101 (Perkin Elmer) and aligned in

Clustal X (Thompson et al 1997) Manual adjustments were made visually by

inserting gaps where necessary

Site Selection

A one-year-old plantation (200 ha E dunnii) approximately 15 km south of Miriam

Vale in southern Queensland was selected as a site to conduct pathogenicity

experiments Very few pests or pathogens were found within the site at the

beginning of the experiment and moderate to low rainfall had occurred during the

previous months No trees were observed to have canker symptoms

An experimental area was selected at the western end of the site which was

relatively flat with clay rich loamy soil The experimental area was surrounded on all

sides by at least 50 m of plantation trees Two experiments were conducted in this

area one inoculated in winter and a second inoculated in summer (100 m apart)

The trees were approximately three metres tall and relatively healthy at the

beginning of the experiment

Cultures and Inoculation

Four-week-old cultures (species to be discussed) grown on half strength PDA were

taken into the field in sealed sterile zip lock bags to prevent contamination The

228

Petri-dishes were handled using latex gloves and were only opened to cut and

remove 5 mm cubes from each culture during inoculation

Inoculation involved cutting a 2 cm wide crescent into the main stem of the tree at a

height of 14 m using a sterile razor blade The depth of the cut was approximately 2

mm deep which exposed the vascular cambium beneath the bark A 5 mm cube of

myceliaagar was placed mycelial surface down beneath the cut wood before

immediately being taped closed with Parafilm SMI tape

Pathogenicity Experiment One (Winter)

The winter pathogenicity experiment was conducted in July 2006 over a six week

period Ten rows of sixteen trees (160 trees) were marked out with flagging tape and

wooden stakes to form a large rectangular block Each tree was then randomly

marked with one of eight different colours of flagging tape to ensure a random

design Each colour of flagging tape was indicative of one of eight pathogenicity

treatments (fungi combinations) (Table 61)

Up to three cubes were placed beneath the bark adjacent to each other (vertically

along the stem) in treatments involving multiple species infections All trees were

inoculated on the same day and were left for 12 weeks before examination

229

Treatment Species combinations

1A Control (sterile agar)

2A Holocryphia eucalypti

3A Neofusicoccum ribis

4A Cytospora eucalypticola

5A H eucalypti + N ribis

6A H eucalypti + C eucalypticola

7A N ribis + C eucalypticola

8A H eucalypt + N ribis + C eucalypticola

Pathogenicity Experiment Two (Summer)

The summer pathogenicity experiment was conducted in November 2006 More

isolates of each pathogen species had been collected prior to the trial and these

were incorporated into the experiment to examine intra-species pathogenicity

Twelve rows of twenty trees (120 trees) were marked out with wooden stakes and

flagging tape adjacent to pathogenicity experiment one (winter) The trees were

randomly assigned to treatments and then inoculated in twelve different

combinations (20 trees treatment) (Table 62)

Table 61 Treatments in the winter pathogenicity experiment 2006 Eight different combinations of inoculations using single isolates of three species of pathogens

230

Treatment Isolate Number amp Species

1B Control

2B 1 Holocryphia eucalypti



5B 1 Neofusicoccum ribis





10B 1 Cytospora eucalypticola




Treatment 1B was a control (water agar) and treatments 2B 5B and 10B used the

same isolates as those used the winter pathogenicity experiment (2A 3A and 4A)

Only these isolates are therefore comparable between the winter and summer

experiments

Measuring Lesions

After 12 weeks the inoculated trees were examined This involved removing the

tape from each stem examining the symptoms and quantifying the damage

Examinations of each lesion involved recording discolouration kino exudation

cracking sinking or swelling and the presence of fruiting structures Quantifying

damage involved measuring the length and width of lesions To increase the

accuracy of measuring lesions the outer layer of bark was first removed by gently

scraping a sterile razor blade over the bark surface The length and width of each

lesion was measured using a 300 mm ruler These two figures were multiplied to

give a Lesion Severity Index (mm)

Table 62 Summer experiment 2006 Twelve individual treatments of different isolates of canker fungi

231

After completing all measurements each lesion was cut from the stem using a sharp

knife These were labelled and refrigerated until they could be further examined This

reduced the likelihood of accidentally introducing pathogens to the plantation and

provided material to conduct Kochrsquos Postulates Kochrsquos Postulates was conducted

using the same methods previously described to isolate and identify pathogens from

naturally occurring basal cankers

Statistics

Lesion Severity Index was used as the response variable Data were analysed using

Statistica (version 6 2004) statistical package For data collected for both

pathogenicity experiments analyses of variance (ANOVA) were carried out for each

treatment

Results

The majority of inoculated trees responded to the pathogens in two ways Trees

either produced a

1) Negative disease response Stems were not infected by pathogens and

produced a light brown callus in response to wounding (Figure 6-2A) or

2) Positive disease response Stems were infected with pathogens and

produced a dark necrotic lesion which often penetrated the bark surface and

was associated with cracking sinking and swelling (Figure 6-2B)

232

Winter Pathogenicity Results

Treatment 1A (control) had an infection rate of 20 This was equal lowest with

treatment 4A (C eucalypticola) and treatment 7A (N ribis + C eucalypticola) (Table

63) The greatest infection rates caused by single species were caused by

treatments 2A (H eucalypti) and 3A (N ribis) which were both 40 The greatest

infection rate caused by a combination of species was caused by treatment 8A (H

eucalypti + N ribis + C eucalypticola) which was 55

Mean Lesion Severity Index was lowest in treatment 1A (control) followed by

treatment 4A (C eucalypticola) (Figure 6-3) The greatest Mean Lesion Severity

Index occurred in treatment 2A (H eucalypti) Significant (Plt005) differences

occurred between treatment 1A (control) and all other treatments between treatment

2A (H eucalypti) and treatment 4A (C eucalypticola) and between treatment 4A (C

eucalypticola) and treatment 8A (H eucalypti + N ribis + C eucalypticola) (Table

Figure 6-2 Two host responses after inoculation with canker pathogens A arrow points to a healed callus response with no resulting infection after inoculation B arrow points to a dark necrotic lesion (infection) with sinking and cracking of the bark (W Lesion width L Lesion length)

W

L

233

64)

Treatments TM1 TM2 t-value df p Valid N Valid N StdDev StdDev F-ratio p

Treatments 1A and 2A 1195 39290 227857 38 0028401 20 20 2524 74726 8764723 0000000






Treatments 2A and 4A 39290 4860 -20393 38 004842 20 20 74726 10814 477515 0000000


Treatment No Canker Fungi Percentage of lesions

1A Control 20

2A H eucalypti 40

3A N ribis 40

4A C eucalypticola 20

5A H eucalypti + N ribis 45

6A H eucalypti + C eucalypticola 35

7A N ribis + C eucalypticola 20

8A H eucalypt + N ribis + C eucalypticola 55

Table 63 Winter pathogenicity experiment Percentage of lesions resulting from 20 stem inoculations for each treatment

Table 64 Winter pathogenicity experiment Analysis of variance (ANOVA) Comparing different treatments (only those which were significant when Plt005 were included) Treatment mean 1 (TM1) Treatment mean 2 (TM2) Degrees of Freedom (df) P value (P) number samples (Valid N) Standard deviation (Std Dev)

00

1000

2000

3000

4000

5000

6000

Control

H eucalypti

B ribis

C eucalyptic

ola

H eucalypti +

B ribis

H eucalypti +

C e

ucalypticola

B ribis + C

euca

lyptic

ola

H eucalypt +

B ribis

+ C e

ucalypticola

Treatments

Mean

Lesio

n S

everi

ty I

nd

ex (

len

gth

x w

idth

) m

m LSD (5) = 12210

Figure 6-3 Winter Pathogenicity Experiment Mean Lesion Severity Index for each treatment Error Bars =SE LSD =Least Significant Difference

1A

2A

3A

4A

5A

6A 7A

8A

234

Summer Pathogenicity Results

The lowest rate of infection was caused by treatment 1B (control) which was 20

(Table 65) The greatest rate of infection was caused by treatment 3B (2 H

eucalypti) which was 100

The lowest Mean Lesion Severity Index was caused by treatment 1B (control)

followed by treatment 8B (4 N ribis) (Figure 6-4) The greatest Mean Lesion

Severity Index was caused by treatments 12B (3 C eucalypticola) and 13B (4 C

eucalypticola) ANOVA showed significant differences (Plt005) in Mean Lesion

Severity Index between treatment 1B (control) and all other treatments (Table 66)

Treatment Canker Fungi Percentage of lesions

1B Control 20

2B 1 H eucalypti 90

3B 2 H eucalypti 100

4B 3 H eucalypti 80

5B 1 N ribis 95

6B 2 N ribis 80

7B 3 N ribis 50

8B 4 N ribis 70

9B 5 N ribis 70

10B 1 C eucalypticola 80




Table 65 Summer pathogenicity experiment Percentage of lesions resulting from 20 stem inoculations in each of thirteen different treatments

235

Treatments TM1 TM2 t-value df p Valid N Valid N G1 StdDev G2 StdDev F-ratio p

Treatments 1B and 2B 880 12620 302 38 00045 20 20 3751 16972 2047 0000000











Winter Versus Summer Pathogenicity

The same number of lesions were caused by treatment 1A (control winter

pathogenicity experiment) and 1B (control summer pathogenicity experiment) (Table

63 and Table 65) Treatment 2A (H eucalypti Winter Pathogenicity Experiment)

caused 40 lesions while treatment 2B (H eucalypti summer pathogenicity

experiment) caused 90 lesions Treatment 3A (N ribis winter pathogenicity

experiment) caused 40 lesions while treatment 5B (1 N ribis summer

Pathogenicity Experiment) caused 95 lesions Treatment 4A (C eucalypticola

Table 66 Analysis of Variance (ANOVA) Summer pathogenicity trial Comparing 13 treatments (Mean lesion severity index) (Only those which were significant (Plt005) are included) Treatment mean 1 (TM1) Treatment mean 2 (TM2) Degrees of Freedom (df) P value (P) number samples (Valid N) Standard deviation (Std Dev)

Figure 6-4 Summer pathogenicity experiment Mean Lesion Severity Index for each treatment (refer to Table 65) Error Bar = SE LSD = Least Significant Difference

0

50

100

150

200

250

300

350

400

450

500

Con

trol

1 H

euca

lypt

i

2 H

euca

lypt

i

3 H

euca

lypt

i

1 B

rib

is

2 B

rib

is

3 B

rib

is

4 B

rib

is

5 B

rib

is

1 C

euca

lypt

icol

a

2 C

euca

lypt

icol

a

3 C

euca

lypt

icol

a

4 C

euca

lypt

icol

a

Isolate species

Lesio

n S

everi

ty I

nd

ex (

len

gth

x w

idth

)

mm

LSD (5) = 7500

1B

2B 3B

4B

5B

6B

7B

8B 9B

10B 11B

12B 13B

236

winter pathogenicity experiment) caused 20 lesions while treatment 10B (1 C

eucalypticola summer pathogenicity experiment) caused 80 lesions

The Mean Lesion Severity Index was similarly low in both treatment 1A treatment

(control winter pathogenicity experiment) and treatment 1B (control summer

pathogenicity experiment) (Figure 6-5) Mean Lesion Severity Index was greater in

treatment 2A (H eucalypti winter pathogenicity experiment) than treatment 2B (H

eucalypti summer pathogenicity experiment) Mean Lesion Severity Index was

greater in treatment 3A (N ribis winter pathogenicity experiment) than treatment 5B

(1 N ribis summer pathogenicity experiment) Mean Lesion Severity Index was

greater in treatment 10B (1 C eucalypticola summer pathogenicity experiment)

than treatment 4A (C eucalypticola winter pathogenicity experiment) (Figure 6-5)

Although Mean Lesion Severity Index varied between the winter and summer

treatments ANOVA showed no significant (Plt005) differences between any

treatments

Fungal Species

Figure 6-5 Winter versus summer pathogenicity Mean Lesion Severity Index for each treatment Error Bars = SE

00

1000

2000

3000

4000

5000

6000

Cont

rol

H e

ucalyp

ti

B r

ibis

C e

ucalyp

ticola

Fungal species

Mean

Lesio

n S

everi

ty I

nd

ex (

len

gth

x

wid

th)

mm

control Holocryphia eucalypti

Neofusicoccum ribis

Cytospora eucalypticola

Winter

Summer

237

Kochrsquos Postulates

As previously described isolation of canker fungi was attempted from diseased stem

tissue which was removed from each of the inoculated trees in both the winter and

summer pathogenicity experiments Fungi were successfully isolated from 95 of all

lesions and 80 of these were a positive match with the species used in the original

inoculation It can therefore be confidently assumed that the majority of the observed

disease symptoms in both experiments were caused by the isolate used in each

treatment

Failure to isolate fungi from 5 of the tissue samples was due to a lack of any fungal

growth in the medium In the remaining 15 of mismatched fungi most of these

samples were contaminated by ubiquitous saprophytes such as Penicillium

Discussion

Both pathogenicity experiments were successful in that all species of canker fungi

caused a disease response in the E dunnii hosts No trees died as a result of the

inoculations but severe infection and potential deaths may have occurred if the

infected material had not been removed from the plantation

Variability in both the percentage of lesions (infections) and the severity of lesions

(Mean Lesion Severity Index) was observed when different pathogens were

inoculated individually and in combination

Pathogenicity between Species

The number of lesions resulting from infection and the Mean Lesion Severity Index

were used as measures of pathogenicity in each of the treatments Significant

differences were observed between the control and all other treatments in the winter

experiment which indicated that trees were responding to inoculation by fungi by

238

producing a disease response However large variability in the disease response

was also observed within treatments (as indicated by large error bars) This

variability indicated that individual trees were responding differently to inoculation by

the same fungal isolates Differences in susceptibility between trees may have been

attributed to genetic differences (Dungey et al 1997) or to differences within the

immediate environment of each tree (Durzan 1974) A lack of rainfall in the

experimental site may also have been a factor Moisture stress can cause greater

susceptibility to canker pathogens (Bertrand et al 1976 Yarwood 1959 Colhoun

1973 Hepting 1963 Boyer 1995 Schoenweiss 1975 1981)

A significant difference in Mean Lesion Severity Index was observed in the winter

pathogenicity experiment between H eucalypti and C eucalypticola H eucalypti

also caused the greatest number of lesions in this experiment which was consistent

with other studies which show that H eucalypti is the most pathogenic of the three

species (Old et al 1990) Given that N ribis C eucalypticola and H eucalypti are all

known to have endophytic characteristics (Bettucci et al 1999 Slippers et al 2004) it

was expected that some trees would not produce a disease response These trees

formed a callus over the wounded area which was also observed by Bettucci and

Alonso (1997) when inoculating seedlings with H eucalypti and C chrysosperma

Unlike the winter pathogenicity experiment no significant differences in pathogenicity

were observed between species in the summer pathogenicity experiment

Pathogenicity within Species

In the summer pathogenicity experiment it was expected that different fungal isolates

of the same species would differ in their pathogenicity (Yuan and Mohammed 2000)

However the only significant difference in Lesion Severity Index occurred between

the control and other treatments

239

Interactions of Pathogens

Three hypotheses were tested which described the interactions of pathogens within

a living host These were







The winter pathogenicity experiment showed that there was a significant difference

in the Mean Lesion Severity Index between treatments 4A (C eucalypticola) and

treatment 8A (H eucalypti + N ribis + C eucalypticola) Given that treatment 8A (H

eucalypti + N ribis + C eucalypticola) caused a greater Mean Lesion Severity Index

than treatment 4A (C eucalypticola) this effect is most consistent with the

Synergism Hypothesis However it must also be recognised that no other treatment

involving inoculation of more than one pathogen produced a significantly greater

disease response It is therefore likely that this difference may be solely due to the

greater pathogenicity of H eucalypti This would support the No Effect Hypothesis

Pathogenicity Summer versus Winter

The climate in southern Queensland is subtropical and it was therefore expected that

trees in the winter pathogenicity experiment would receive less rain than those in the

summer pathogenicity experiment It was also expected that the trees would become

stressed during periods of low rainfall which would lead to increased susceptibility to

pathogens However the summer of 2006 received lower than average rainfall which

240

meant that the summer climate was similar to the winter climate at least in terms of

rainfall

No significant differences in Mean Lesion Severity Index were observed between the

shared isolates of the winter and summer pathogenicity experiments This was

mainly due to lsquobackground noisersquo caused by large variability within each treatment A

greater number of lesions occurred in the summer experiment than the winter

experiment when comparing treatments 2A and 3B (H eucalypti) and treatments 3A

and 5B (N ribis) however the opposite effect occurred when comparing treatments

4A and 10B (C eucalypticola) A contradiction in the winter versus summer effect

also occurred due to the generally greater number of lesions observed in summer

and the generally greater Mean Lesion Severity observed in winter

Conclusion

The main finding of the study was that H eucalypti appears to be more pathogenic

than C eucalypticola (based on Mean Lesion Severity Index) and N ribis (based on

number of lesions) However due to confounding factors such as atypical climate a

controlled glasshouse experiment may have been more informative Repeating the

experiment during more typical climatic conditions in the absence of drought may

also yield better results

It is interesting that a disease response occurred in some of the control treatments of

both the winter and summer pathogenicity experiments Given that the wounded

stems were sealed with tape these lesions may have been caused by latent

pathogens already occurring within the stems Isolation of fungi from these lesions

revealed the presence of other species such as the saprophytes Cladosporium spp

and Alternaria spp These species are not considered pathogenic and it is therefore

241

possible that host stress (caused by drought conditions) may have made the trees

more susceptible to infection by common saprophytes

242

7 General Discussion

Important Pests

Important pests were identified during the survey and included defoliators

sapsuckers and borers Although most of these species caused low levels of

damage some species caused severe damage which could potentially lead to

economic loss The two most important pests identified were Paropsisterna cloelia

(Chrysomelidae) and Endoxyla cinerea (Cossidae)

P cloelia was by far the most abundant defoliating insect species in southern

Queensland The beetles prefer to feed on young expanding foliage (de Little and

Madden 1975 Tanton and Khan 1978) and the regrowth produced by drought

stressed plantation trees appeared to benefit their development Greater levels of

damage were observed in the northern plantations which was probably due to the

greater quantities of foliar regrowth being produced in this region (due to higher

annual rainfall) The northern plantations also appeared to have greater vigour in

terms of general tree health and a generally greater rate of recovery after defoliation

Borer species were less common in plantations compared to defoliator species The

most abundant borer species was the giant wood moth E cinerea The larvae of

these moths caused severe localised damage in several plantations Unlike

defoliators which generally cause severe damage due to their high numbers wood

moths may cause severe damage as individuals A single larva can compromise the

health of a plantation tree by causing stem breakage Wood moths preferred to

attack E grandis rather than E dunnii however concern about impacts has been

reduced due to the growing trend toward planting E dunnii

243

Important Pathogens

The number of foliar pathogens identified during the study was lower than expected

Dry conditions can have adverse affects on pathogen lifecycles by reducing

sporulation and spore dispersal (Howe 1955 Walklate et al 1989 Daniel and Shen

1991 Agrios 2005) These conditions can also reduce the infection success of fungal

spores (Beaumont 1947 Krausse and Massie 1975) It is hypothesised that the

drought in eastern Australia created adverse conditions for the development of many

foliar pathogens during the study period

Despite the dry conditions a number of important pathogens were identified in the

genera Teratosphaeria and Mycosphaerella Teratosphaeria spp are common in

eucalypt plantations throughout Australia and new species are often described

(Crous 1998) Several species are common in Queensland (Sivanesan and Shivas

2002) including T cryptica (Park and Keane 1982a Park and Keane 1982b Crous

and Wingfield 1996 Park et al 2000) For unknown reasons T cryptica was never

found during the study Severe outbreaks of Mycosphaerella heimii were identified

This species was previously known only from Madagascar and Indonesia (Whyte et

al 2005)

The most common canker pathogen was H eucalypti This pathogen was identified

as the causal agent of lsquosudden death syndromersquo H eucalypti was the only pathogen

found to cause tree deaths in plantations Given that the disease became less

common as the study progressed this indicates that the fungus is adversely affected

by drought conditions Consequently it is recommended that plantation health

surveys continue to monitor this pathogen in the future (especially during years of

high rainfall)

244

Economic Impacts

It is important to note that damage caused by pests and diseases can only cause

economic impacts in eucalypt plantations if stem growth and wood volume are

compromised or if trees are killed (Judd 1996) Angel et al (1999) estimated that

stem growth of E dunnii is adversely affected when crown damage exceeds 375

Other eucalypt species have similar thresholds (Elek 1997) During the present

study chrysomelid damage reached a maximum of 519 and it is therefore likely

that economic impacts occurred E cinerea (giant wood moth) and H eucalypti

(canker pathogen) also caused economic impacts by causing localised tree deaths in

some plantations It is recommended that further research examines similar damage

thresholds for other important pests and diseases

Pest and Pathogen Management

Current pest management in southern Queensland involves the use of generalist

insecticides Although these insecticides have beneficial short-term lsquoknock downrsquo

effects it is likely that more indirect processes will ultimately contribute to long-term

pest management For example the negative impacts of herbivory may be greatly

reduced by increasing the vigour of plantation trees (Stone 1991) This may be

achieved through better species site selection (Howe 1955 Stork 1988 Hill 1994

Nair 2001) or through selectively breeding for greater resistance (Painter 1951

Maxwell and Jennings 1980 McDonald 1981 Eldridge et al 1994 Barbour 1997

Soria and Borralho 1998 Jones et al 2002) Other methods of reducing impacts of

pests in plantations include reducing transmission between plantations (Floyd et al

1998) or by reducing monoculture effects (Root 1973)

Despite the impacts of foliar pathogens and canker pathogens no control methods

for diseases are currently employed in southern Queensland Given that chemical

245

control is usually inefficient (Dickman 1992) the most common method of controlling

pathogens is to selectively breed plantation trees with greater resistance (Alfenas et

al 1983 Dianese et al 1984 Ostry and McNabb 1986 Denison and Kietzka 1993

Alfenas et al 1997 van Heerden and Wingfield 2002 Gryzenhout et al 2003) Given

that H eucalypti was the only pathogen observed to be capable of causing tree

deaths selectively breeding for resistance to cankers may be a viable option for the

future

General Plantation Health

Several studies show that pest and pathogen species cannot be viewed in isolation

without considering the interaction and the extrinsic influence of environmental

factors (Coley et al 1985 Waring and Cobb 1992 Rand 1999 Straus and Agrawal

1999) Effectively every interaction between a host pest or pathogen is an

interaction of the hostrsquos genotype the pest or pathogenrsquos genotype and the

environment (Matheson and Cotterill 1990 Basford and Cooper 1998) Although a

number of important factors have been identified as influencing plantation health in

the present study additional factors should also be considered The following factors

were identified as having increasingly adverse effects in plantations which were

exposed to drought conditions

1 Soil Characteristics On several occasions it was observed that severely

moisture stressed plantations occurred on shallow soils Studies show that

shallow soils have a lower capacity to store water (Aspinall 1965 Bachelard

1986) Low soil water storage capacity can also exacerbate the impacts of

drought

2 Topography and Aspect Trees on north facing slopes appeared be exposed

246

to higher temperatures and therefore suffered greater moisture stress than

those on more protected south facing slopes (Fekedulegn et al 2003) Trees

on the crests of hills where lsquorun offrsquo was greatest were also more prone to

moisture stress Healthier trees generally occurred at the base of large slopes

and within drainage lines

3 Weeds Plantations with high infestations of weeds appeared to suffer due to

greater competition with weeds for water and nutrients Previous studies show

that moisture stress in E dunnii plantations can be reduced through weed

control and fertiliser application (Stone and Birk 2001 Xu and Dell 1997 Xu et

al 2002)

4 Tree Density Tree growth may have been compromised due to over stocking

which caused competition between individual trees Competition appeared to

be greatest in areas with low soil fertility or greater weather exposure (crests

north facing slopes)

Tree Decline Conceptual Models

Manionrsquos Tree Decline Spiral

The lsquoTree Decline Spiralrsquo created by Manion (1981) is a model which illustrates how

various negative impacts (both abiotic and biotic) may contribute to the decline of

tree health (Figure 7-1) The many interacting factors driving the decline spiral are

divided into predisposing inciting and contributing factors Predisposing factors are

the background abiotic components of a particular environment and the unique

properties of the trees therein On the other hand the inciting and contributing

factors are mainly the background of biotic stress agents Severe episodic lsquoacts of

godrsquo such as frost drought or human-caused stresses are also included among the

247

inciting factors

Figure 7-1 The tree decline spiral by Manion (1981) illustrating the range of factors which may contribute to ultimate tree death

Modified Tree Decline Spiral

A modified spiral has been developed to illustrate how some of the more important

abiotic and biotic factors may contribute to tree deaths in eucalypt plantations in

southern Queensland (Figure 7-2) Factors feeding into the outer spiral are mainly

those which should be reduced or controlled before a plantation is established

Factors on the inner spiral are those which tend to have influences after plantations

have been established Smaller secondary spirals for defoliating insects and insect

borers have also been included to illustrate the feedback loop mechanisms which

can occur when these insects repeatedly attack severely stressed trees (Carne

248

1965 Landsberg 1990a Landsberg 1990b Landsberg 1990c Stone and Bacon


Figure 7-2 A modified tree decline spiral to illustrate some of the more important factors which may contribute to tree death in plantations

The lsquoTree Decline Spiralrsquo is particularly useful in showing the range of factors which

may contribute to tree death however the structure of the spiral is limited in that it

suggests that the processes of tree decline only goes in one direction The

implication is that the potential for recovery of a stressed tree is not suitably

illustrated

The Tree Recovery Decline Seesaw

The findings of the present study show that stressed trees in plantations can recover

rapidly following adequate rainfall Trees can alternate between various stages of

health (indefinitely) before succumbing to death depending on the degree of stress

249

caused by various adverse biotic and abiotic factors A new model has therefore

been developed to illustrate the processes related to both tree decline and recovery

The lsquoTree RecoveryndashDecline Seesawrsquo illustrates how drought stressed trees can

either respond to recovery after rainfall or may continue to decline during prolonged

drought conditions The model also illustrates five stages of varying tree health and

the processes by which pests and diseases may exploit stressed trees (Figure 7-3)

Figure 7-3 The lsquoTree Recovery Decline Seesawrsquo which illustrates the opposing outcomes of recovery and decline of a stressed tree in response to impacts by pests and diseases and the influence of rainfall and drought

250

The lsquoTree RecoveryndashDecline Seesawrsquo is a unique approach to illustrating the

process of tree decline and recovery Support for each stage of the model is evident

in both the findings of this study and the scientific literature (Tables 71 amp 72)

251

Process of Tree Decline

Stage 1 Stage -1 Stage -2

Tree Health Pest Impacts Pathogen Impacts Tree Health Pest Impacts Pathogen Impacts Tree Health Pest Impacts Pathogen Impacts

Tree is moderately drought stressed and shows a deterioration of foliage including leaf necrosis (Stone and Bacon 1994 Landsberg 1990)

Borers are attracted to the moderately stressed tree (Lieutier 2002)

This supports the plant stress hypothesis (Koricheva and Larsson 1998)

Latent pathogens initiate a disease response within the tissues of the moderately drought stressed tree (Anselmi et al 2007)

Tree is severely drought stressed and is suffering from leaf necrosis and premature leaf loss (Stone and Bacon 1994 Landsberg 1990)

Defoliating insects are attracted to the severely stressed tree (White 1969 Larsson and BjOumlrkman 1993) This supports the plant stress hypothesis

Primary pathogens fail to sporulate or spread via splash dispersal due to dry conditions (Howe 1955 Walklate et al 1989 Daniel and Shen 1991 Agrios 2005)

Tree has succumbed to death due to prolonged drought stress

Defoliating insects are disadvantaged by poor host quality (Rouault et al 2006)

Saprophytic fungi infect dead tissue (Bier 1959 1961 Griffin 1977 Rayner and Boddy 1988 Bettucci and Saravay 1993 Bettucci and Alonso 1997 Bettucci et al 1999)

Table 71 The lsquoTree Recovery-Decline Seesawrsquo illustrating the outcome of decline of a stressed tree in response to impacts by pests and diseases and the influence of drought Relevant literature is sited for individual impacts of pests and pathogens

Threshhold

Pivot Threshhold

Pivot

Threshhold Pivot

252

Process of Tree Recovery

Stage 1 Stage +1 Stage +2

Tree Health Pest Impacts

Pathogen Impacts Tree Health Pest Impacts

Pathogen Impacts Tree Health Pest Impacts Pathogen Impacts





Tree is in a state of recovery and as such it is producing flush regrowth (Jacobs 1955)

Defoliating insects such as chrysomelid beetles are attracted to the new foliar regrowth (Tanton and Khan 1978 Edwards 1982 Edwards and Wanjura 1990)

This supports the plant vigour hypothesis (Price 1991)

Latent pathogens resume a latent state within the tissues of the recovering tree (Anselmi et al 2007)

Tree is in an optimum state of health where growth and recovery are at a maximum

Pest impacts are ameliorated by host vigour (Benson and Hager 1993 Stone 2001)

Pathogen impacts are ameliorated by host vigour (Benson and Hager 1993 Stone 2001)

Table 72 The lsquoTree Recovery-Decline Seesawrsquo illustrating the outcome of recovery of a stressed tree in response to impacts by pests and diseases and the influence of rainfall Relevant literature is sited for individual impacts of pests and pathogens

Threshhold Pivot

Threshhold Pivot

Threshhold Pivot

253

Mechanisms of the lsquoTree Recovery-Decline Seesawrsquo are consistent with hypotheses

and explanations related to the influence of drought stress on pests and diseases (as

discussed in previous chapters) Some of the more important features of the model are

as follows

1 Processes of tree decline illustrate the processes of the lsquoPlant Stress Hypothesisrsquo

(White 1969 1984 Louda and Collinge 1992 Koricheva and Larsson 1998)

2 Processes of tree recovery illustrate the processes of the lsquoPlant Vigour

Hypothesisrsquo (Price 1991 Inbar et al 2001)

3 Levels of host stress (moderate to severe) have differing effects on insect

feeding guilds (foliar pests and borers)

4 Levels of host stress (moderate to severe) have differing effects on fungal guilds

(latent pathogens primary pathogens and saprophytic fungi)

5 Stressed trees may alternate (indefinitely) between recovery and decline before

eventually succumbing to death This is also consistent with the description of a

lsquoFeedback Loop Mechanismrsquo (Carne 1965 Landsberg 1990a Landsberg 1990b

Landsberg 1990c Stone and Bacon 1995 Landsberg and Cork 1997)

Although the lsquoTree Recovery Decline Seesawrsquo is a simplistic model it does provide a

unique approach to understanding the complex interactions of stressed trees and

their associated pests and pathogens Given that each stage of the model presents

a unique set of abiotic and biotic conditions the model also provides an explanation

for why pests and pathogens are diverse in plantations The changing conditions

within each stage provide a mode of niche partitioning which is implicated as an

important factor for the co-occurrence of species (Schoener 1974) There is also

254

potential for application of the model to other abiotic factors which influence pests

and pathogen susceptibility such as stress caused by waterlogged soil or nutrient

deficiency

Limitations of the Study

The following limitations of the study were identified

1 Drought Impacts

The effects of drought on plantation trees and their associated pests and pathogens

was ecologically interesting however it was unfortunate that the initial aims of the

study were largely jeopardised due to drought effects Factors which were expected

to influence the abundance of pests and pathogens such as plantation age local

climate season and tree species were largely overshadowed by the effects of

drought This was mainly due to the rapid rates of leaf loss and regeneration which

occurred in moisture stressed trees

Interpreting the results of experiments was difficult because variation in levels of

damage could be equally attributed to either the effects of leaf loss or regeneration

The drought was therefore an unfortunate confounding factor

2 Sampling Methodology

The modified version of the Crown Damage Index Assessment (CDIA) was used to

assess the percentage of damage within tree canopies for different types of damage

(Stone et al 2003) This sampling methodology was limited because it did not

account for leaf loss or regeneration between samples Experimental error may have

occurred due to inaccurate estimates in damage levels which may also have been

accelerated by the drought conditions Over-estimating levels of damage in

255

plantations has occurred in similar studies Reichle et al (1973) estimated that levels

of damage were over-estimated by 65 Abbott et al (1993) estimated a

comparable over-estimate of 57

One method of reducing experimental error would be to count all the damaged

leaves on a set number of branches on one side of the tree Ohmart et al (1985)

suggested counting damaged leaves as a way of assessing insect damage but

emphasised the limitations of the technique over a long time period because leaves

may be removed by mechanisms other than feeding insects Lowman (1984)

suggests a similar methodology but also suggested that the assessment should be

restricted to expanding leaves

Sampling could be further improved by assessing levels of damage at different

levels within the tree canopy Ohmart et al (1983a) showed that the lower crowns of

eucalypts are often more heavily defoliated than the upper crowns Observations of

defoliating species such as chrysomelid beetles would suggest that the opposite

effect occurs in E dunnii plantations in southern Queensland This information may

have been valuable given that vertical tree growth is largely dependent on growth at

the apex of the crown and that damage in this area may have a more negative effect

on tree growth than damage at the base of the crown (Ohmart et al 1983a)

Other methodologies for assessing the size of pest populations may involve the

identification of symptoms other than leaf damage Defoliating insect species

sometimes produce faeces which are identifiable to species level (Jacobs 1955)

The amount of faeces in leaf litter can be correlated with visual ratings of insect

defoliation (Edwards et al 1993 Pook et al 1998)

256

3 Specimen collection

The majority of samples of diseased material were collected from either foliage or

stems of plantation trees Very few samples were collected from root tissue This

was mainly because in the few instances where roots were excavated they

appeared to be healthy with no disease symptoms However more sampling may

have revealed a greater diversity of root borne pathogens

4 Sampling Regime

Observations in the field showed that sudden changes in the abundance of pests

and pathogens could occur within a week or even a few days Such changes were

often triggered by rainfall events after prolonged dry periods Because sampling

occurred every three months it is likely that variability in the abundance of pests and

pathogens may have been missed between samples Financial constraint due to the

large distance between the university in Perth and the study sites in southern

Queensland was a limiting factor Assessing the abundance of damage more

frequently during the year and correlating this with more detailed rainfall history may

have improved the study

5 Weather Data

Rainfall in southern Queensland was found to be sporadic and highly variable over

short distances (2-3km) Weather data was used from weather stations (BOM)

occurring approximately 10 km from plantations being assessed Given this

distance data could only be used as a rough guide as to the amount of rainfall

actually received by plantations In hindsight the use of individual weather stations

within each plantation would have made examining the effects of climate and season

257

more efficient

Future Research

Although a large diversity of pests and pathogens were examined during the course of

the study no single species were examined in great detail A better understanding of

the biology of individual species is essential to developing species specific control

methods Future research is recommended for the following species

1 Chrysomelid Beetles

P cloelia is an important pest of plantations in southern Queensland A number of

colour forms of the species were identified which suggested a species complex may

occur Taxonomic research of P cloelia at both a morphological and molecular level

would help determine the relatedness of these different colour forms If the species

was found to be a species complex this raises the question as to whether all

species are as destructive as each other in plantations This may lead to targeted

surveys within plantations and an examination of the life history characteristics of

each potential species Important pest characteristics may include fecundity the rate

of larval development and an examination of the feeding capacities of both larvae

and adults

Developing control methods for P cloelia may include testing the effects of various

insecticides on beetles in controlled experiments Other important research may

include identifying the over-wintering sites of adult beetles to allow pest control

during the winter months Identifying the natural predators of species may allow the

development of a biological control agent (Baker et al 2003)

258

2 Leaf Pathogens

Teratosphaeria spp and Mycosphaerella spp are abundant in Australian eucalypt

plantations Despite the relatively small number of species collected during the

study it remains likely that a greater diversity of undescribed species are yet to be

identified in southern Queensland Further sampling during summer periods is likely

to produce more new species

3 Canker Pathogens

H eucalypti was identified as being the only pathogen capable of killing its host The

species is therefore the greatest disease threat to the plantation industry in southern

Queensland Although H eucalypti was identified as the causal pathogen of lsquoSudden

Death Syndromersquo the cause of the apparent random incidence of this disease is yet

to be determined Also it is yet to be determined whether the species is introduced

to plantations with seedlings or associated soil A targeted survey for the disease in

native forest may help elucidate its origin

Concluding Remarks

Despite the hindrance of the drought and the shortcomings of some of the experimental

designs the findings of the study provide valuable insight into the role of drought stress

in plantations and its various effects on pests and diseases Little research has been

conducted to date for many of these species and a more detailed understanding is

required if the plantation industry in southern Queensland is to reach its full potential

259

8 References

ABARE 2009 The Australian Bureau of Agricultural and Resource Economics

Australian Forest and Wood Product Statistics March and June Quarters 2009

Abbott I Smith R Williams M and Voutier R 1991 Infestation of regenerated

stands of karri (Eucalyptus diversicolor) by bullseyes borer (Tryphocaria

acanthocera Cerambycidae) in Western Australia Australian Forestry 54 66-

74

Abbott I 1991 Insect pest problems of eucalypt plantations in Australia 6 Western

Australia Australian Forestry Journal 56 381-384

Abbott I Van Heurck P Burbridge T and Williams M 1993 Damage caused by

insects and fungi to eucalypt foliage spatial and temporal patterns in

Mediterranean forest of Western Australia Forest Ecology and Management 58

85-110

Abebe G and Hart JH 1990 The relationship of site factors to the incidence of

Cytospora and Septoria cankers and poplar and willow borers in hybrid poplar

plantation United States Department of Agriculture and Forestry Services

Technical Report NC-272

Adams AJ Wingfield MJ Common R and Roux J 2005 Phylogenetic

relationships and morphology of Cytospora species and related teleomorphs

(Ascomyces Diaporthales Vasaceae) from Eucalyptus Studies in Mycology 52

1-44

Adams MA and Atkinson PI 1991 Nitrogen supply and insect herbivory in

eucalypts the role of nitrogen assimilation and transport processes In

lsquoProductivity in Perspectiversquo (eds PJ Ryan) pp 239-241 Third Australian Forest

Soils and Nutrition Conference Melbourne October 1991 Forestry Comission of

New South Wales Sydney

Agosteo GE Pennisi A M 1990 Discovery of Hypoxylon mediterraneum on chestnut

in Calabria Journal Tecnica Agricola 42(1) 55-59

260

Agrios GN 1980 Insect involvement in the transmission of fungal pathogens In

lsquoVectors of Plant Pathogensrsquo (Eds FK Harris and K Maramorosch) pp 234-293

Academic Press New York

Agrios G N 2005 Plant Pathology Fifth Edition Academic Press

Alfaro RI Omule S A Y 1990 The effect of spracing on Sitka Spruce Weevil

damage to Sitka Spruce Canadian Journal of Forestry Research 20 179-184

Alfenas AC Jeng R and Hubbes M 1983 Virulence of Cryphonecria cubensis on

Eucalyptus species differing in resistance European Journal of Forest Pathology

13 197-205

Alfenas AC Valle LAC Xavier AA Brommonschenkel SH Grattapaglia D

Silva CC Bertolucci FL and Penchel R 1997 Eucalyptus rust genetic

variability of elite clones and histological characterization of the resistance

reaction In lsquoProc IUFRO Conf on Silviculture and Improvement of Eucalypt Vol

2rsquo pp 60ndash64 Salvador Bahia Brazil

Altieri MA and Letourneau DK 1984 Vegetation diversity and insect pest outbreaks

CRC A Critical Review Plant Science 2 131-169

Anagnostakis SL 1984 The effect of temperature on growth of Endothia

(Cryphonectria) parasitica in vitro and in vivo Mycologia76(3) 387-397

Anderson AB 1990 Steps towards sustainable use of the Amazon Rain Forest In

lsquoAlternatives to Deforestationrsquo Columbia University Press New York

Andow DA 1991 Vegetational diversity and arthropod population response Annual

Review of Entomology 26 561-586

Andjic V Barber PA Carnegie AJ Hardy GEStJ Wingfield MJ and Burgess

TI 2007 Phylogenetic reassessment supports accommodation of

Phaeophleospora and Colletogloeopsis from eucalypts in Kirramyces

Mycological Research 111(10) 1184-1198

Andjic V Barber PA Carnegie AJ Pegg GS Hardy GEStJ Wingfield MJ

and Burgess TI 2007 Kirramyces viscidus sp nov a new eucalypt pathogen

from tropical Australia closely related to the serious leaf pathogen Kirramyces

destructans Australasian Plant Pathology 36 478ndash487

261

Angel PJ Nichols JD and Stone C 1999 Growth increments of Eucalyptus dunnii

subsequent to damage by Creiis lituratus (Hemiptera Psyllidae) Proceedings

2003 ANZIF Conference Queenstown New Zealand

Anilla E 1969 Influence of temperature upon the development and voltinism of Ips

typhographus L (Coleoptera Scolytidae) Annual Review of Entomology 6 161-

167

Anselmi N Nasini M Mazzaglia A Librandi A Rocco E Ravaioli F 2007

Correlation between the occurrence of pathogenic fungal endophytes in healthy

oak trees and oak decline Journal of Plant Pathology 89 (3) 28

Appel DN and Stipes RJ 1986 A description of declining and blighted pin oaks in

eastern Virginia Journal of Arboriculture 12 155-158

Arnold AE Maynard Z and Gilbert GS 2000 Are tropical fungal endophytes

hyperdiverse Ecological Letters 3 267-274

Arnold RJ Gardiner G Wang G Zhang J and Wu Z 1998 Genetic variation and

selection of Eucalyptus dunnii in China New Forests 19(3) 215-226

Aspinall D 1965 The effects of soil moisture stress on the growth of barley Australian

Journal of Agricultural Research 16 265-275

Attiwill PM 1994 Ecological disturbance and the conservative management of

eucalypt forests in Australia Forest Ecology and Management 63 (2-3) 301-

346

Avtzis DN Arthofer W Stauffer C Avtzis N Wegensteiner R 2010 Pityogenes

chalcographus (Coleoptera Scolytinae) at the southernmost borderline of

Norway spruce (Picea abies) in Greece Entomologia Hellinica 19 3-13

Bachelard EP 1986 Effects of soil moisture stress on growth of seedlings of three

eucalypt species III Tissue-water relations Australian Forestry Research 16

155-163

Bailey C and Duncanson T 1998 From blue sky to blue chip Landscope 14 35-42

Baker SA Elek JA Bashford R Paterson SC Madden J and Battaglia M 2003

Inundative release of Coccinellid beetles into eucalypt plantations for biological

control of chrysomelid leaf beetles Agricultural and Forest Entomology 5(2) 97ndash

106

262

Ball MC Egerton JJG Leuning R and Cunninham RB 1997 Microclimate abobe

grass adversely affects spring growth of seedlings snowgum (Eucalyptus

pauciflora) Plant Cell Environment 20 155-166

Bamber RK Humphreys FR 1965 Variation in sapwood starch levels in some

Australian forest species Australian Forestry 2 15-23

Barbour L 1997 Breeding better blue gums Landscope 13 36-41

Barker J L 1979 Geographical variations in spore morphology of Diplodia pinea For

Comm Wood Tecchnology Division New South Wales Sydney

Barr ME 1972 Preliminary studies on the Dothideales in temperate North America

Contributions from the University of Michigan Herbarium 9 523-638

Basford KE and Cooper M 1998 Genotype x environment interactions and some

considerations for wheat breeding in Australia Australian Journal of Agricultural

Research 49 153-174

Bauhus J Khanna PK and Menden N 2000 Aboveground and belowground

interactions in mixed plantations of Eucalyptus globulus and Acacia mearnsii

Canadian Journal of Forestry 30(12) 1886-1894

Beadle CL 2000 Physiology of eucalypts in relation to disease In lsquoDiseases and

Pathogens of Eucalyptsrsquo (eds PJ Keane G A Kile F D Podger and B N

Brown) pp 61-68 CSIRO Publishing Melbourne

Beadle CL and Inions G 1990 Limits to growth of Eucalyptus and their biology of

production In lsquoProspects for Australian Plantationsrsquo (eds J Dargavel and N

Semple) pp 183-193 Centre for Resource and Environmental studies Australian

National University Canberra

Beadle CL Turnbull CRA and Dean GH 1996 Environmental effects on growth

and kraft pulp yield of Eucalyptus globulus and Eucalyptus nitens Appita Journal

49 239-42

Beaumont A 1947 The dependence on the weather of the dates of potato blight

epidemics Transactions of the British Mycological Society 31 45-53

Bell DT 1999 Australian trees for the rehabilitation of waterlogged and salinity

damaged landscapes Australian Journal of Botany 47 697-716

263

Benson JS and Hager TG 1993 The distribution abundance and habitat of

Eucalyptus dunnii (Myrtaceae) (Dunrsquos white gum) in New South Wales

Cunninghamiana 3(1) 123-144

Bertrand PF English H Uria K and Schick FJ 1967 Late season water deficits

and development of Cytospora canker in French prune Phytopathology 66

1318-1320

Bertus AL and Walker J 1974 Ramularia on Eucalyptus and Angophora

Australasian Plant Pathology Society Newsletter 3 3

Bettucci L and Alonso R 1997 A comparative study of fungal populations in healthy

and symptomatic twigs of Eucalyptus grandis in Uruguay Mycological Research

101 1060-1064

Bettucci L Alonso R and Tiscornia S 1999 Endophytic mycobiota of healthy twigs

and the assemblage of species associated with twig lesions of Eucalyptus

globulus and E grandis in Uruguay Mycological Research 103(4) 468-472

Bettucci L and Saravay M 1993 Endophytic fungi of Eucalyptus globulus a

preliminary study Mycological Research 97(6) 679-692

Bier JE 1939 Septoria canker of native and introduced hybrid poplars Canadian

Journal of Forestry Research 17 195-204

Bier JE 1959 The relation of bark moisture to the development of canker diseases

caused by native facultative parasites I Cryptodiaporthe canker on Willow

Canadian Journal of Botany 37 229-238

Bier JE 1961 The relation of bark moisture to the development of canker disease

casued by native facultative parasites VI Pathogenicity studies of Hypoxylon

pruinatum (Klotzsch) and Septoria musiva on species of Acer Populus and Salix


Birch TTC 1937 Diplodia pinea in New Zealand Review of Applied Ecology 16 148

Blake T BevilacquaE Barbosa MDM 1990 In lsquoEarly Selection of Fast-Growing

Eucalyptus Clones and Speciesrsquo pp 26-34 IPEF International Piracicaba

Boland DJ Brooker MIH Chippendale CM Hall N Hyland PBM Johnston

R Kleinig DA and Turner JD 1984 In lsquoForest Trees of Australiarsquo pp 687

Nelson and CSIRO Melbourne

264

Booth TH Stein JA Nix HA and Hutchinson MF 1989 Mapping regions

climatically suitable for particular species An example using Africa Forest

Ecology and Management 28 19-31

Booth TH and Jones PG 1998 Identifying climatically suited areas for growing

particular trees in Latin America Forest Ecology and Management 108 167-

173

Boyer JS 1995 Biochemical and Biophysical aspects of water deficits and the

predisposition to disease Annual Review of Phytopathology 33 251-274

Bren LJ and Gibbs NL 1986 Relationships between flood frequency vegetation

and topography in a river red gum forest Australian Forest Research 16 357-

370

Bradford KJ and Hsiao TC 1982 Stomatal behaviour and water relations of

waterlogged tomato plants Plant Physiology 70 1508-1513

Brennan EB and Weinbaum SA 2001 Effect of epicuticular wax on adhesion of

psyllids to glaucous juvenile and glossy adult leaves of Eucalyptus globulus

Labillardiere Australian Journal of Entomology 40 270-277

Brennan EB Weinbaum SA Rosenheim JA and Karban R 2001 Heteroblasty in

Eucalyptus globulus (Myricales Myricaceae) Affects ovipositonal and settling

preferences of Ctenarytaina eucalypti and C spatulata (Homoptera Psyllidae)

Environmental Entomology 1 1144ndash1149

Brewer R and Merritt P G 1978 Wind throw and tree replacement in a climax

beech-maple forest Oikos 30 (1) 149-152

Briggs JD and Leigh JH 1988 In lsquoRare or Threatened Australian Plantsrsquo pp 278

Australian national parks and wildlife service special publication No 14 ANPWS

Canberra

Britton E B 1970 Coleoptera In lsquoThe Insects of Australiarsquo A Textbook for Students

and Research Workers and the Supplement (ed Waterhouse DF) First Edition

Melbourne University Press

Brown BN Bevege DI and Steven RE 1981 Site stress and Diplodia induced

dieback and death of hail damaged slash pine XVII IUFRO Congress Kyoto

Japan

265

Brooker I and Kleinig DA 1990 In lsquoA Field Guide to Eucalypts South Eastern

Australia Vol 1rsquo Blooming Books Australia

Brooker MIH and Slee AV 1996 Dicotyledons Winteraceae to Myrtaceae In lsquoFlora

of Victoria Vol 3rsquo (eds Walsh NG and Entwisle TJ) Inkata Press Melbourne

Bruck RI and Manion PD 1980 Interacting environmental factors associated with

the incidence of Hypoxolon canker on trembling aspen Canadian Journal of

Forestry Research 10 17-24

Buchanan TS 1967 Diplodia twig blight of Pine In lsquoImportant Forest Insects and

Diseases of Mutual Concern to Canada United states and Mexicorsquo pp 189-191

North American Forestry Commission Canadian Department of Forestry

Ottawa

Buffo E Battisti A Stastny M and Larsson S 2007 Temperature as a predictor of

survival of the pine processionary moth in the Italian Alps Agricultural and Forest

Entomology 9 65-72

Burgess TI and Wingfield MJ 2002 Impact of fungi in natural forest ecosystems A

focus on eucalypts In lsquoMicrorganisms in Plant Conservation and Biodiversityrsquo

(eds K Sivasithamparam KW Dixon and RL Barret) pp 285-306 Kluwer

Academic Publishers Dordrecht

Burgess TI Gordon TR Wingfield MJ and Wingfield BD 2004 Geographic

isolation of Diplodia scrobiculata and its association with native Pinus radiata

Mycological Research 108 1399-1406

Burgess TI Sakalidis ML and Hardy GEStJ 2006 Gene flow of the canker

pathogen Botryosphaeria australis between Eucalyptus globulus plantations and

native eucalypt forests in Western Australia Austral Ecology 31 559-566

Burns RM and Honkala BH 1990 Silvics of North America Vol 2 Hardwoods

Agriculture Handbook 654 US Department of Agriculture Forest Service

Washington DC

CALM 1990 Calm Insect Manual CALM Perth Western Australia

Cameron PJ Hill RL Bain J and Thomas WP 1993 Analysis of importations for

biological control of insect pests and weeds in New Zealand Biological Science

and Technology 3 387-404

266

Campbell KG 1972 Insect hazards in monoculture tree plantations as opposed to

mixed planting and natural forest associations in Australia Forestry Log 5 42-

44

Candy SG Elliot HJ Bashford R Greener A 1992 Modelling the impact of

defoliation by the leaf beetle Paropsisterna bimaculata (Coleoptera

Chrysomelidae) on height growth of Eucalyptus regnans Forest Ecology and

Management 54 69-87

Carne PB 1965 Distrabution of the eucalyptus feeding sawfly Perga afinis Australian

Journal of Ecology 13 593-612

Carne PB 1966 Ecological characteristics of the eucalypt-defoliating chrysomelid

Paropsis atomaria Australian Journal of Zoology 14 647-672

Carne PB Greaves TG and McInnes RS 1974 Insect damage to plantation-grown

eucalypts in north coastal New South Wales with particular reference to

Christmas beetles (Coleoptera Scarabaeidae) Journal of the Australian

Entomological Society 13 189-206

Carnegie AJ 2000 A study of the species of Mycosphaerella species on eucalypts

and the impacts of Mycosphaerella species on Eucalyptus globulus Labill PhD

Thesis University of Melbourne

Carnegie AJ 2002 In lsquoA Field guide to Common Pests and Diseases in Eucalypt

Plantations in NSWrsquo (State Forests of New South Wales Sydney)

Carnegie AJ 2007a Forest health condition in New South Wales Australia 1996-

2005 I Fungi recorded from eucalypt plantations during forest health surveys

Australasian Plant Pathology 36 213-224

Carnegie AJ 2007b Forest health condition in New South Wales Australia 1996-

2005 II Fungal damage recorded from eucalypt plantations during forest health

surveys and their managment Australasian Plant Pathology 36 1-15

Carnegie AJ Ades PK Keane PJ and Smith IW 1994 Provenance variation in

Eucalyptus globulus in susceptibility to Mycosphaerella leaf disease Canadian


267

Carnegie AJ Ades PK Keane PJ and Smith IW 1998 Mycosphaerella diseases

of juvenile foliage in a eucalypt species and provenance trial in Victoria

Australia Australian Forestry 61 190-194

Carnegie AJ and Ades PK 2000 The proportion of leaf spots caused by

Mycosphaerella cryptica and M nubilosa on Eucalyptus globulus E nitens and

their F1 hybrids in a family trial in Tasmania Australia Australasian Mycologist

21 (2) 53-63

Carnegie AJ and Ades PK 2003 Mycosphaerella leaf disease reduces growth of

plantation grown Eucalyptus globulus Australian Forestry 66 113-119

Carnegie AJ and Keane PJ 1994 Further Mycosphaerella species associated with

leaf diseases of Eucalyptus Mycological Research 98 413-418

Carnegie AJ Keane PJ and Podger FD 1997 The impact of three species of

Mycosphaerella newly recorded on Eucalyptus in Western Australia Australasian

Plant Pathology 26 (2) 71-77

Carnegie AJ and Keane PK 2002 The proportion of leaf spots caused by

Mycosphaerella cryptica and Mycosphaerella nubilosa on Eucalyptus globulus

E nitens and their F1 hybrids in a family trial in Tasmania Australia Australasian

Mycology 21(2) 53-63

Carnegie A J and Angel P 2005 Creiis lituratus (Froggatt) (Hemiptera Psyllidae) a

new insect pest of Eucalyptus dunnii plantations in sub-tropical Australia

Australian Forestry 68(1) 59-64

Carnegie AJ Stone C Lawson SA and Matsuki M 2005b Can we grow certified

eucalypt plantations in subtropical Australia ndash an insect pest management

perspective New Zealand Journal of Forestry Science 35 223-245

Carroll FE Muller E and Sutton BC 1977 Preliminary studies on the incidence of

needle endophytes in some European conifers Sydowia 29 87-103

Carroll GC 1988 Fungal endophytes in stems and leaves From latent pathogens to

mutualistic symbiont Ecology 69 2-9

Carroll GC and Carroll FE 1978 Studies on the incidence of Coniferous needle

endophytes in the pacific north west Canadian Journal of Botany 56 3034-3040

268

Carter W 1973 In lsquoInsects in Relation to Plant Disease 2nd edrsquo John Wiley and Sons

London

Cesati V and De Notaris G 1963 Schema di classificazione deglisferiacei italici

aschigeri piu o meno appartenenti al genere Sphaeria nellrsquoantico significato

attribuitoglide Persoon Comment Soc Crittog Ital 1(4) 177-240

Chauzat MP Purvis G and Dunne R 1995 Release and establishment of a

biological control agent Psyllaephagus pilosus for eucalyptus psyllid

(Ctenarytaina eucalypti) in Ireland Annals of Applied Ecology 141(3) 293-304

Cheah LH 1977 Aerobiology and epidemiology of Mycosphaerella nubilosa (Cke)

Hansf on Eucalyptus spp MSc University of Auckland New Zealand

Chipompha NWS 1987 Phaeoseptoria eucalypti a new pathogen of Eucalyptus in

Malawi South African Forestry Journal 142 10-12

Chippendale GM 1988 Myrtaceae Eucalyptus Angophora In lsquoFlora of Australia Vol

19rsquo Australian Government Publishing Services Canberra

Chou CKS 1982 Susceptibility of Pinus radiata seedlings to infection by Diplodia

pinea as affected by pre-innoculation conditions New Zealand Journal of Forest

Sciences 12 438-441

Christiansen CM 1940 Studies on the biology of Valsa saldida and Cytospora

chrysosperma Phytopathology 30 459-475

Christiansen E 1992 After-effects of drought did not predispose young Picea abies to

infection by the bark beetle-transmitted blue-stain fungus Ophiostoma polonicum

Scandinavian Journal of Forestry Research 7 557-569

Christiansen E and A Bakke 1988 The spruce bark beetle of Eurasia In lsquoDynamics

of Forest Insect Populationsrsquo (eds A A Berryma) pp 479-503 Plenum

Publishing Corporation New York amp London

Christiansen E Bakke A 1997 In lsquoProceedings Integrating cultural tactics into the

management of bark beetle and reforestation pestsrsquo USDA Forest Service

General Technical Report NE-236

Christiansen E and A Ericsson 1986 Starch reserves in Picea abies in relation to

defence reaction against a bark beetle transmitted blue-stain fungus

Ceratocystis polonica Canadian Journal of Forestry Research 16 78-83

269

Clark LR 1962 The general biology of Cardiaspina albitextura (Psyllidae) and its

abundance in relation to weather and parasitism Australian Journal of Zoology

10 537-586

Clarke KR amp Gorley RN 2001 PRIMER v5 userrsquos manual tutorial PRIMER-E Ltd

Plymouth UK

Coley PD Bryant JP and Chapin FS 1985 Resource availability and plant

antiherbivore defence Science 230 895-899

Collet N 2001 Insect pests of young eucalypt plantations Agricultural Notes AG0799

ISSN 1329-8061 Forest Science centre Heidelberg

Colhoun J 1973 Effects of environmental factors on plant disease Annual Review of

Phytopathology 11 343-364

Common IFB 1970 Lepidoptera In lsquoThe Insects of Australiarsquo A Textbook for

Students and Research Workers and the Supplement (ed Waterhouse DF)

First Edition Melbourne University Press

Common IFB 1990 In lsquoMoths of Australiarsquo Melbourne University Press Melbourne

Cook RJ and Papendick RI 1972 Influence of water potential of soils and plants on

root diseases Annual Review of Phytopathology 10 349-374

Cooper PD 2001 What physiological processes permit insects to eat eucalypt leaves

Austral Ecology 26 556-562

Corlett M 1995 An annotated list of the published names in Mycosphaerella and

Sphaerella Corrections and Additions Mycotaxon 53 37-56

Cortinas MN Burgess TI Dell B Xu D Crous PW Wingfield B and Wingfield

M J 2006 First record of Colletogloeopsis zuluense comb nov causing a stem

canker of Eucalyptus in China Mycological research 110 229-236

Cotterill PP Moran GF and Grigg BR (1985) Early growth of 36 species of

eucalypts near Mount Gambier South Australia Australasian Forestry Research

15 409-416

County P and County N 2003 Wood decay fungi in landscape trees In lsquoPest Notes

no 74109rsquo

270

Crist CR Shoeneweiss DF 1975 The influence of controlled stresses on

susceptibility of European White Birch stems to attack by Botryosphaeria

dothidea Phytopathology 65 369-373

Croiseacute L and Lieutier F1993 Effects of drought on the induced defence reaction of

Scots pine to bark beetle associated fungi Annals of Forestry Science 50 91-

97

Cromer RN and Williams ER 1982 Biomass and nutrient accumulation in a planted

Eucalyptus globulus (Labill) fertiliser trial Australian Journal of Botany 30 265-

278

Crous PW 1998 In lsquoMycosphaerella spp and their Anamorphs Associated with Leaf

Spot Diseases of Eucalyptusrsquo The American Phytopathological Society St

Paul Minnesota USA

Crous PW Slippers B Wingfield MJ Rheeder J Marasas WFO Philips AJL

Alves A Burgess TI Barber PA and Groenewald JZ 1996 Phylogenetic

lineages in the Botryosphaeriaceae Studies in Mycology 55(1) 235-253

Crous PW Knox-Davies PS and MJ Wingfield 1989a Infection studies with

Phaeoseptoria eucalypti and Coniothyrium ovatum on Eucalyptus spp South

African Forestry Journal 149 30-35

Crous PW Knox-Davies PS and Wingfield MJ 1989b A summary of fungal leaf

pathogens of Eucalyptus and the diseases they cause in South Africa South


Crous PW and Wingfield MJ 1996 Species of Mycosphaerella and their anamorphs

associated with leaf blotch disease of eucalypts in South Africa Mycologia 88

441-458

Crous PW Braun U Groenewald JZ 2007 Mycosphaerella is polyphyletic Studies in

Mycology 58 1ndash32

Crous PW Wingfield MJ Mohammed C Yuan and Z Quing 1998 New foliar

pathogens from Australia and Indonesia Mycological Research 102 (5) 527-

532

Curry SJ 1981b The association of insects with eucalypt dieback in southwestern

Australia In lsquoEucalypt Dieback in Forests and Woodlandsrsquo (eds KM Old GA

271

Kile CP Ohmart) CSIRO Melbourne

Daniel WS and Shen KW 1991 Threshold relative humidity forecasts for plant

disease prediction Journal of Applied Meteorology 30 463-477

Davidson C Gottschalk K Johnson J 1999 Tree Mortality Following Defoliation by

the European Gypsy Moth (Lymantria dispar L) in the United States A Review

Forest Science 45 (1) 74-84

Davies J M and King C J 1977 Pine Shoot Beetles Forestry Commission Leaflet 3

HMSO London

Davison EM 1982 Endothia havanensis on Jarrah Australasian Plant Pathology 11

10-11

Davison EM and Coates DJ 1991 Identification of Cryphonectria cubensis and

Endothia gyrosa from eucalypts in Western Australia using isozyme analysis


Davison EM and Tay CS 1983 Twig branch and upper trunk cankers of Eucalyptus

marginata Plant Disease 67 1285-1287

Day JS 1998 Light conditions and the evolution of heteroblasty (and the divaricate

form) in New Zealand New Zealand Journal of Ecology 22 43-54

de Little DW and Madden JL 1975 Host preference in the Tasmanian eucalypt

defoliating Paropsini (Coleoptera Chrysomelidae) with particular reference to

Chrysoptharta bimaculata (Olivier) and C agricola (Chapuis) Journal of the

Australian Entomolgical Society 14 387-294

de Little DW 1989 Paropsine chrysomelid attack on plantations of Eucalyptus nitens

in Tasmania New Zealand Journal of Forestry Science 19 223-227

Dell B and Malajczuk N 1994 Boron deficiency in eucalypt plantations in China

Canadian Journal of Forestry Research 24 2409-2416

Denison NP and Kietzka JE 1993 The development and utilisation of vegetative

propagation in Mondi for commercial afforestation programmes South African

Bosboutydskr 165 47-54

272

Desprez-Loustau ML Marccedilais B Nageleisen LM Piou D Vannini A 2006

Interactive effects of drought and pathogens in forest trees Annals of Forest

Science 63 597-612

DeWitt JR and Ambrust EJ 1978 Feeding preference studies of adult Nezara

viridula (Linnaeus) (Hemiptera Pentatomidae) morphs from India and the United

States Great Lakes Entomology 11(1) 67-69

DeWitt NB and Godfrey GL 1972 A bibliography of the southern green stink bug

Nezara viridula (Linnaeus) (Hemiptera Pentatomidae) ILL Natural History Survey

Biological Notes 78 23

Dianese JC Moraesde TSA and Silva AR 1984 Response of Eucalyptus species

to field infection by Puccinia psidii Plant Disease 68 314-316

Dick M 1982 Leaf-inhabiting fungi of eucalypts in New Zealand New Zealand Journal

of Forestry Science 12 525-527

Dickman A 1992 Plant Pathogens and Long-Term Ecosystem Changes In lsquoThe

Fungal Community Its Organisation and Role in the Ecosystemrsquo (eds GC Caroll

amp DT Wicklow) New York

Dobbertin M Wermelinger B Bigler C Buumlrgi M Carron M Forster B Gimmi U

Rigling A 2007 Linking Increasing Drought Stress to Scots Pine Mortality and

Bark Beetle Infestations The Scientific World Journal 7(1) 231-239

Drake CJ 1920 The southern green stink bug in Florida Florida State Plant Board Q

Bulletin 4 41-94

Duffy EAJ 1963 A monograph of the immature stages of Australasian timber beetles

British Museum of Natural History London pp 235

Dungey HS Potts BM Carnegie AJ and Ades PK 1997 Mycosphaerella leaf

disease genetic variation in damage to Eucalyptus nitens E globulus and their

F1 hybrid Canadian Journal of Forestry Research 27 250-259

Dungey HS Potts BM Whitham TG and Li H 2000 Plant genetic affects

community richness and composition Evidence from a synthetic eucalypt hybrid

population Evolution 54 1938-1946

Duniway JM Gordon TR 1986 Water relations and pathogen activity in soil Journal

of Experimental Botany 35 1782-1786

273

Durzan DJ 1974 In lsquoNutrition and Water Relations of Forest Trees a Biochemical

Approachrsquo pp 15-63 Proceedings Third North American Forest Biology

Workshop

Eastwood R 2004 Successive replacement of tending ant species at aggregations of

scale insects (Hemiptera Margarodidae and Eriococcidae) on Eucalyptus in

south-east Queensland Australian Journal of Entomology 43(1) 1ndash4

Edmunds GF Jr and Alstad DN 1978 Coevolution in insect herbivores and conifers

Science 199 941-945

Edwards PB 1982 Do waxes on juvenile eucalypt leaves provide protection from

grazing insects Australian Journal of Ecology 7 347-352

Edwards PB and Wanjura WJ 1990 Physical attributes of eucalypt leaves and the

host range of chrysomelid beetles Synopsia Biologica Hungarica 39 227-236

Edwards PB Wanjura WJ and Brown WV 1993 Selective herbivory by Christmas

beetles in response to intraspecific variation in Eucalyptus terpenoids Oecologia

95 551-557

Eldridge KG 1961 Significance of Diplodia pinea in Plantations Review of Applied

Mycology 41 339-339

Eldridge K Davidson J Harwood C and van Wyk G 1994 In lsquoEucalypt

Domestication and Breedingrsquo pp 288 Clarendon Press Oxford

Eldridge K Kent DS and Urquhart CAP 1995 The control of insects on eucalypts

Forest Protection Research Division Series No C1 State Forests of New South

Wales Beecroft NSW

Elek JA 1997 Assessing the impact of leaf beetles in eucalypt plantations and

exploring options for their management Tasforests 9 139-153

Elliot H J Bashford R Greener A and Candy SG 1992 Integrated pest

management of the Tasmanian Eucalyptus leaf beetle Chrysophtharta

bimaculata (Olivier) (Coleoptera Chrysomelidae) Forest Ecology and

Management 53 (1-4) 29-38

Elliot HJ and de Little DW 1985 In lsquoInsect Pests of Trees and Timber in Tasmaniarsquo

Forestry Commission of Tasmania Hobart

274

Elliot HJ Kile GA Candy SG and Ratkowsky DA1987 The incidence and spatial

pattern of Nothofagus cunninghamii (Hook) Oerst attacked by Platypus

subgranosus Schedl in Tasmanias cool temperate rainforest Australian Journal

of Ecology 12(2) 125-138

Elliott HJ and Greener A 1994 Prospects for using Bacillus thuringiensis for control

of eucalypt leaf beetles in Tasmania In lsquoProceeding of the second Canberra

Bacillus thuringiensis meeting 21-23 September 1993rsquo (ed RJ Akhurst) pp 147-

151 CSIRO Division of Entomology Canberra

Elliot HJ Ohmart CP and Wylie FR 1998 In lsquoInsect pests of Australian forests

Ecology and Managementrsquo pp 214 Inkata press Melbourne

Eyles AC 1999 Introduced Mirinae of New Zealand (Hemiptera Miridae) New

Zealand Journal of Zoology 26 355-372

FAO (Food and Agriculture Organisation for the United Nations) 1988 The Eucalypt

Dilemma FAO Rome

FAO Global forest resources assessment 2010- Main report FAO Forestry paper

(available at httpwwwfaoorgforestryfoframainindexjsp)

Farr JD 2002 Biology of the gumleaf skeletoniser Uraba lugens Walker (Lepidoptera

Noctuidae) in the southern jarrah forest of Western Australia Australian Journal

of Entomology 41(1) 60ndash69

Farrell GS and New TR 1980 Some aspects of the biology of the eucalypt sawfly

Phylacteophaga froggati Riek (Hymenoptera Pergidae) Australian Journal of

Zoology 28 83-90

Farrell RCC Bell DT Akilan K and Marshall JK 1996 Morphological and

Physiological Comparisons of Clonal Lines of Eucalyptus camaldulensis I

Responses to Drought and Waterlogging Australian Journal of Plant Physiology

23(4) 497-507

Faulds W 1991 Spread of Bracon phylacteophagus a biocontrol agent of

Phylacteophaga froggatti and impact on host New Zealand Journal of Forestry

Science 21 185-193

275

Fekedulegn D Hicks RR and Colbert JJ 2003 Influence of topographic aspect

precipitation and drought on radial growth of four major tree species in an

Appalachian watershed Forest Ecology and Management 177 (1-3) 409-425

Ferreira CA do Couto HTZ and Van Wyk G 1993 The influence of environmental

variables on the growth of speciesprovenances of Eucalyptus species in the

states of Minas Gerais and Espirito Santo Boletim de Persquisa Florestal 3 9-

25

Ferreira FA and Milani D 2002 lsquoVisual Diagnosis and control of abiotic and biotic

Eucalyptus diseases in Brazilrsquo (International Paper Sacirco Paulo Brasil)

Fisher PJ and Petrini O 1990 A comparative study of fungal endophytes in leaves

xylem and bark of Almas species in England and Switzerland Mycological

Research 94 313-319

Fisher PJ and Petrini O 1992 Fungal saprobes and pathogens as endophytes of

rice (Oryza sativa L) New Phytologist 120 137-143

Fisher PJ Petrini O and Sutton BC 1993 A comparative study of fungal

endophytes in leaves xylem and bark of Eucalyptus nitens in Australia and

England Sydowia 45 1-14

Flanagan JG 1994 The Australian distribution of Mictis profana (F) (Hemiptera

Coreidae) and its life cycle on Mimosa pigra Australian Journal of Entomology

33 (2) 111ndash114

Fletcher MJ 1985 Revision of the genus Siphanta Staringl (Homoptera Fulgoroidea

Flatidae) Australian Journal of Zoology Supplementary 33(110) 1 ndash 94

Fletcher MJ 2008 Illustrated Key to the Genera of the family Aphrophoridae

found in Australia (Hemiptera Cercopoidea) Online Document 7243 Orange

Agricultual Instituite NSW Department of Primary Industries

Flock RA 1957 Biological notes on a new Chalcid-fly from seed-like Eucalyptus galls

in California Pan-Pacific Entomologist 33 153-155

Florence RG 1996 Ecology and Silviculture of Eucalypt Forests CSIRO Australia

276

Floyd R Wylie R Old K Dudzinski M and Kile G 1998 Pest risk analysis of

Eucalyptus spp at risk from incursions of plant pests and pathogens through

Australiarsquos northern border CSIRO Contracted Report No 44 CSIRO

Publishing

Fox LR and Macauley BJ 1977 Insect grazing on Eucalyptus in response to

variation in leaf tannins nitrogen Oecologia 29 145-162

Fox LR and Morrow PA 1983 Estimates of damage by herbivorous insects on

eucalyptus trees Australian Journal of Ecology 8 139-147

Franceschini A Linaldeddu BT Pisanu P Pisanu S 2004 Effects of water stress

on the endophytic incidence of Biscogniauxia mediterbanea in cork oak trees

Journal of Plant Pathology 86(4) 319-320

Fraser D and Davison EM 1985 Stem cankers of Eucalyptus saligna in Western


Fry G 1983 Eucalyptus in New Zealand A position report New Zealand Journal of

Forestry 28 394-411

Gardes M and Bruns T 1993 ITS primers with enhanced specificity for

basidiomycetes ndash application to the identification of Mycorrhizae and rusts

Molecular Ecology 2 113-118

Gasow H 1925 Der gr une Eichenwickler als Forstsch adling Arbeiten aus der

biologischen Reichsanstalt fur Land- und Forstwirtschaft 12 355-508

Gavran M and Parsons M 2011 Australian plantation statistics 2011 Australian

Bureau of Agricultural and Resource Economics and Sciences Canberra

Grezahgne A Cortinas MN Wingfield MJ and Roux J 2005 Characterisation of

the Coniothyrium stem canker pathogen on Eucalyptus camaldulensis in

Ethiopia Australasian Plant Pathology 34 1-6

Gibbs JN 1997 Fifty years of sooty bark disease of sycamore Quarterly Journal of

Forestry 91 215-221

Gibson IAS 1975 Diseases of forest trees widely planted as exotics in the tropics and

southern hemisphere Part 1 Important members of the Myrtaceae

Leguminosae Verbinaceae and meliaceae Commonwealth Mycological Institute

and Forestry Institute Kew and Oxford

277

Gibson IAS 1980 Two pine needle fungi new to Columbia Tropical Pest

Management 26 38-40

Goodyer GJ 1985 Chinese junk caterpillars Agfact AE36 Department of

Agriculture NSW Agdex 622

Graham RD and Webb MJ 1991 Micronutrients and resitance and tolerance in

plants In lsquoMicronutrients in Agriculture 2nd editionrsquo pp 329-370 SSSA Book

series No 4

Griffin DM 1977 Water potential and wood decay fungi Annual Review of


Griffiths M Wylie R Lawson S Pegg G and McDonald J 2004 Known or

potential threats from pests and diseases to prospective tree species for high

value timer plantings in northern Australia Mareeba Department of Primary

Industries and Fisheries Horticulture and Forestry science Indooroopilly

Queensland 4068 Australia

Grundy P and Maelzer D 2000 Assessment of Pristhesancus plagipennis (Walker)

(Hemiptera Reduviidae) as an augmented biological control in cotton and

soybean crops Australian Journal of Entomology 39 (4) 305-309

Gryzenhout M 2006 Microthia Holocryphia and Ursicollum three new species on

Eucalyptus and Cocoloba for fungi previously known as Cryphonectria Studies in

Mycology 55 35-52

Gryzenhout M Eisenberg BE Coutinho TA Wingfield BD and Wingfield MJ

2003 Pathogenicity of Cryphonectria eucalypti to Eucalyptus clones in South

Africa Forest Ecology and Management 176 427-437

Gryzenhout M Myburg H Hodges CS Wingfield BD and Wingfield MJ 2006

Microthia Holocryphia and Ursicollum three new genera of Eucalyptus and

Cocolaba for fungi previously known as Cryphonectria Studies in Mycology 55

35-52

Gueacuterarda N Dreyerb E Lieutiera F 2000 Interactions between Scots pine Ips

acuminatus (Gyll) and Ophiostoma brunneo-ciliatum (Math) estimation of the

critical thresholds of attack and inoculation densities and effects on hydraulic

properties in the stem Annals of Forestry Science 57 681ndash690

278

Guyon JC Jacobi WR and McIntyre GA 1996 Effects of environmental stress on

the development of Cytospora canker of Aspen Plant Disease 80 1320-1326

Hadlington P 1996 Gum tree defoliation by cup moth caterpillars Forest Timber 4(2)

10-11

Hagen KS 1962 Biology and ecology of predacious Coccinellidae Annual Review of

Entomology 7 289-326

Hammer LG Nicholls N Mitchell CD 2000 In lsquoApplications of Seasonal

Forecasting in Agricultural and Natural Systemsrsquo Kluwer Academic Publishers

The Netherlands

Hanks LM Gould JR Pain TD Millar JG and Wang Q 1995 Biology and host

relations of Avetianella longoi (Hymenoptera Encyrtidae) an egg parasitoid of

the eucalyptus longhorned borer (Coleoptera Cerambycidae) Annals of the

Entomological Society of America 88 666-671

Hanks LM Paine TD Millar JG Campbell CD and Schuch UK 1999 Water

relations of host trees and resistance to the phloem-boring beetle Phoracantha

semipunctata F (Coleoptera Cerambycidae) Oecologia 119(3) 400-407

Hansen EM 1999 Disease and diversity in forest ecosystems Australasian Plant

Pathology 28 313-319

Harden GJ 1991 In lsquoFlora of New South Wales Vol 2rsquo UNSW Press Kensington

Harrington RA and Ewel JJ 1997 Invasion of plantations by native and non-

indigenous plant species in Hawaii Forest Ecology and Management 99(1-2)

153-162

Harris VE Todd IW 1980 Temporal and numerical pattern of reproductive

behaviour in the southern green stink bug Nezara Viridula (Hemiptera

Pentatomidae) Entomological Expertise and Application 27(2) 105-116

Hatcher PE 1995 Three way interactions between plant pathogenic fungi herbivorous

insects and their plant hosts Biological Review 70 639-694

Heath RN Roux J Gryzenhout M Carnegie AJ Smith IW and Wingfield MJ

2007 Holocryphia eucalypti on Tibouchina urvilleana in Australia Australasian

Plant Pathology 36 560-564

279

Heather NW 1975 Life history and biology of the leaf bagworm Hyalarcta huebneri

(Westwood) (Lepidoptera Psychidae) Australian Journal of Entomology 14(4)

353ndash361

Heather WA 1967 Susceptibility of the juvenile leaves of Eucalyptus bicostata Maiden

to infection by Phaeoseptoria eucalypti (Hansf) Walker Australian Journal of

Biological Sciences 20 769-775

Hendry SJ Lonsdale D Boddy L 1998 Strip cankering of beech (Fagus sylvatica)

pathology and distribution of symptomatic trees New Phytopathology 140 549-

565

Hepting GH 1963 Climate and forest diseases Annual Review of Phytopathology

1 31-50

Hepting GH 1974 Death of the American chestnut Journal of Forest History 18 60-

67

Hickman GW and Perry EJ 1997 In lsquoTen Common Wood Decay Fungi on

Landscape Trees Identification handbookrsquo Sacramento Western Chapter ISA

Hickman GW and Perry EJ 2003 Wood Decay Fungi in Landscape Trees

Publication Number 74109 In lsquoPest Notesrsquo University of California Cooperative

Extension Nevada

Hill DS 1994 Insect distributions and ecology In lsquoAgricultural Entomologyrsquo Timber

Press Inc Hong Kong

Hillis WE and Brown AG 1987 In lsquoEucalypts for Wood Productionrsquo pp 424 CSIRO

Melbourne

Hillis DM and Huelsenbeck P 1992 Signal Noise and Reliability in Molecular

Phylogenetic Analyses The Journal of Heredity 83(3) 189-19

Hodar JA and Zamora R 2002 Host utilisation by moth and larval survival of pine

processionary caterpillar Thaumetopoea pityocampa in relation to food quality in

three Pinus species Ecological Entomology 27 291-301

Howe RW 1955 The effect of temperature and humidity on the rate of development

and mortality of Tribolium castaneum (Herbst) (Coleoptera Tenebrionidae)

Annals of Applied Ecology 44(2) 356-368

280

Huber JT and Prinsloo GL (1990) Redescription of Anaphes nitens (Girault) and

descriptions of two new species Haliday (Hymenoptera Mymaridae) parasites of

Gonipterus scuttelatus Gylenhall (Coleoptera Curculionidae) in Tasmania

Journal of Australian Entomological Society 29 333-341

Huber JT Mendel Z Protasov A and La Salle J 2006 Two new Australian species

of Stethynium (Hymenoptera Mymaridae) larval parasitoids of Ophelimus

maskelli (Ashmead) (Hymenoptera Eulophidae) on Eucalyptus Journal of

Natural History 40(32) 1909-1921

Huberty A Denno R 2004 Plant water stress and its consequences for herbivorous

insects A new synthsesis Ecology 85 1383-1393

Huelsenbeck JP Bull JJ and Cunningham CW 1996 Combining data in

phylogenetic analysis Trends in Ecology amp Evolution 11(4) 152-158

Hunter GC Crous PW Carnegie AJ Burgess TI and Wingfield MJ 2011

Mycosphaerella and Teratosphaeria diseases of Eucalyptus easily confused and

with serious consequences Fungal Diversity DOI 101007s13225-011-0131-z

Inbar M Doostdar H and Mayer RT 2001 Suitability of stressed and vigorous

plants to various insect herbivores Oikos 94(2) 228-235

Jackson S Maxwell A Neumeister-Kemp HG Dell B and Hardy GEStJ 2004

Infection hyperparasitism and conidiogenesis of Mycosphaerella lateralis on

Eucalyptus grandis in Western Australia Australasian Plant Pathology 33 49-

53

Jackson SL Maxwell A Dell B and Hardy GEStJ 2005 New records of

Mycosphaerella leaf diseases from Eucalypts in Western Australia Australasian


Jacobi WR Riffle JW 1989 Effects of water stress on Thyronectria canker of

Honeylocusts Phytopathology 79 1333-1337

Jacobs MR 1955 Growth habits of the eucalypts Government Printer Canberra

ACT

James DG 1994 Prey consumption by Pristhesancus plagipennis Walker (Hemiptera

Reduviidae) during development Australian Entomologist 21(2) 43-48

281

Jayasinghe CK Silva WPK and Nishantha N 2009 Occurence of Cylindrocladium

quinqueseptatum leaf spot on Hevea brasiliensis in Sri Lanka Journal of

Biological Science 38 (1) 27-30

Jǿker D 2004 Eucalyptus urophylla S T Blake Seed leaflet no 89 Collaboration of

Forest and Landscape and Indonesia Forest Seed Project Horsholm Denmark

Jones T and Gibson IAS 1966 The present world situation in regard to the spread

of internationally dangerous forest diseases and insects In lsquoProcceedings of the

6th World Forestry Congress 2rsquo pp 1897-909

Jones J R 1985 The Distribution of Aspen In lsquoAspen Ecology and Management in

the Western United Statesrsquo (eds N V Debyle and R P Winokur) pp 9-10 U S

Department of Agricultural Forestry Services Rocky Mt Technical Report RM-

119

Jones T H Potts B M Vaillancourt R E and Davies N W 2002 Genetic

resistance of Eucalyptus globulus to autumn gum moth defoliation and the role of

cuticular waxes Canadian Journal of Forestry Research 32(11) 1961-1969

Journet ARP 1980 Intraspecific variation in food plant favourability to phytophagous

insects psyllids on Eucalyptus blakelyi M Ecological Entomology 5 249-261

Jovanovic T Arnold J and Booth T 2000 Determining the climatic suitability of

Eucalyptus dunnii for plantations in Australia China and Central and South

America Journal New Forests 19(3) 215-226

Judd TS 1996 In lsquoNutrition of Eucalyptsrsquo (eds PM Attiwill and MA Adams) pp 249-

258 CSIRO Australia

Kavanagh RP and Lambert MJ 1990 Food selection by the Greater Glider

Petauroides volans is foliar nitrogen a determinant of habitat quality Australian

Wilderness Research 17 285-299

Keane PJ Kile GA Podger FD and Brown BN 2000 In lsquoDiseases and

Pathogens of Eucalyptsrsquo CSIRO Publishing Melbourne Australia

Keane RM and Crawley MJ 2002 Exotic plant invasions and the enemy release

hypothesis Trends in Ecology amp Evolution 17(4) 164-170

Keen NT 1990 Gene-for-gene complementarity in plant-pathogen interactions

Annual Review of Genetics 24 447-463

282

Kendrick B 1992 Fungal Plant Pathology in Agriculture and Forestry Inrsquo The Fifth

Kingdomrsquo pp 193-212 Mycologue Publications

Key KHL 1970 Mantodea In lsquoThe Insects of Australiarsquo (ed DF Waterhouse) pp 294-

301 Melbourne University Press Australia

Khanna PK 1997 Comparison of growth and nutrition of young monocultures and

mixed stands of Eucalyptus globulus and Acacia mearnsii Forest Ecology and

Management 94 105-113

Kile GA 1974 Insect defoliation in the eucalypt regrowth forests of southern

Tasmania Australian Forestry Research 6 9-18

Kile GA and Walker J 1987 Chalara australis sp nov (Hyphomycetes) a vascular

pathogen of Nothofagus cunninghamii (Fagaceae) in Australia and its

relationship to other Chalara species Australian Journal of Botany 35(1) 1-32

Kiritani K Sasaba T 1969 The differences in bio- and ecological characteristics

between neighbouring populations in the southern green stink bug Nezara

viridula Japanese Journal of Ecology 19(5) 177-184

Kirisits T 2004 Fungal associates of European bark beetles with special emphasis on

the ophiostomatoid fungi In lsquoBark and Wood Boring Insects in Living Trees in

Europe a Synthesisrsquo (eds Lieutier F KR Day A Battisti JC Greacutegoire and H

Evans) pp 185-223 pp Kluwer Academic Publishers Dordrecht The

Netherlands

Kolattukudy PE 1985 Enzymatic penetration of the plant cuticle by fungal pathogens

Annual Review of Phytopathology 23 223-250

Kolattukudy PE and Koller W 1983 Fungal penetration of the first line defensive

barriers of plants In lsquoBichemical Plant Pathologyrsquo (eds Wiley) pp 79-100 New

York

Koricheva J and Larsson SH 1998 Insect performance on experimentally stressed

woody plants a meta-analysis Annual Review of Entomology 43 195-216

Kramer PJ 1969 In lsquoPlant and Soil Relationships A Modern Synthesisrsquo pp 482

McGraw and Hill New York

283

Krauss A 1969 Einfluss der Ernahrug der Pflanzen mit mineralstoffen auf den befall

mit parasitaren Krankheiten und Schadlingen Z Pflanzenernahr Bodenkd 124

129-147

Krausse RA and Massie LB 1975 Predictive systems Modern approaches to

disease control Annual review of Phytopathology 13 31-47

Landsberg JJ 1990a Dieback of rural eucalypts Does insect herbivory relate to

dietary quality of tree foliage Australian Journal of Ecology 15 73-87

Landsberg JJ 1990b Dieback of rural eucalypts Response of foliar dietary quality and

herbivory to defoliation Australian Journal of Ecology 15 89-96

Landsberg JJ 1990c Dieback of rural eucalypts The effect of stress on the nutritional

quality of foliage Australian Journal of Ecology 15 97-107

Landsberg JJ and Cork SJ 1997 Herbivory Interactions between eucalypts and the

vertebrates and invertebrates that feed on them In lsquoEucalypt Ecology Individuals

to Ecosystemsrsquo (eds JE Williams JCZ Woinarski) pp 342-372 Australia

Landsberg JJ and Gillieson DS 1995 Regional and local variation in insect

herbivory vegetations and soils of eucalypt associations in contrasted landscape

positions along a climatic gradient Australian Journal of Ecology 20 299-315

Lanfranco D and Dungey HS 2001 Insect damage in Eucalyptus A review of

plantations in Chile Austral Ecology 26 477-481

Laranjeiro AJ 1994 Integrated pest management at Aracruz Cellulose Forest


Larsson S 1989 Stressful times for the plant-stress performance hypothesis Oikos 56

277-83

Larsson S Ekbom B and Bjorkman C 2000 Influence of plant quality on pine saw

fly population dynamics Oikos 89(3) 440-450

Larsson S and Ohmart CP 1988 Leaf age and larval performance of the leaf beetle

Paropsis atomaria Ecological Entomology 13 19-24

Laughton EM 1937 The incidence of fungal disease on timber trees in South Africa

South African Journal of Science 33 377-382

284

Lavallee R 1994 The effects of water stress on the behaviour and development of the

White Pine Weevil Pissodes strobi (Peck) (Coleoptera Curculionidae) on White

Pine Pinus strobes PhD Thesis Concordia University Quebec Canada

Lawrence R Potts BM and Whitham TG 2003 Relative importance of plant

ontogeny host genetic variation and leaf age for a common herbivore Ecological

society of America 84(5) 1171-1178

Lawson SA Wylie FR Wylie RL and Ryan P 2002 Longicorn beetles

(Phoracantha spp) and giant wood moths (Endoxyla spp) emerging threats in

subtropical and tropical eucalypt plantations in Queensland Australia FORSPA

Publication 302002 pp 30-45

Lawton JH 1983 Plant architecture and the diversity of phytophagous insects Annual


Lee DJ Debuse VJ and Pomroy PC 2000 Eucalypt hybrids for commercial farm

forestry in South-East Queensland Final Report National Heritage Trust project

No 982727 pp 28-38

Levitt J 1980 Responses of Plants to Environmental Stresses pp 697 New York and

London Academic

Lichtenthaler HK 1996 Vegetation stress An introduction to the stress concepts in

plants Journal of Plant Physiology 148 4-14

Lieutier F 2002 In lsquoMechanisms of resistance in conifers and bark beetle attack

stategiesrsquo Kluwer Academic Publishers Dordrecht

Lieutier F 2004 In lsquoHost resistance to bark beetles and its variationsrsquo Kluwer


Linnard W 1969 Cultivation of eucalypts in the USSR Forest Abstracts 30 199-209

Loch AD and Floyd RB 2001 Insect pests of Tasmanian blue gum Eucalyptus

globulus globulus in south-western Australia History current perspectives and

future prospects Austral Ecology 26 458-466

Loch AD 2005 Mortality and recovery of eucalypt beetle pests and beneficial

arthropod populations after commercial application of the insecticide a-

cypermethrin Forest Ecology and Management 217 255-265

285

Loch AD 2006 Phenology of Eucalyptus weevil Gonipterus scutellatus Gyllenhal

(Coleoptera Curculionidae) and chrysomelid beetles in Eucalyptus globulus

plantations in south-western Australia Agriculture and Forest Entomology 8(2)

165-185

Loch A D Matthiessen JN Floyd RD 2004 Parasitism and seasonal phenology of

leafblister sawfly Phylacteophaga froggatti (Hymenoptera Pergidae) in

Eucalyptus globulus plantations in south-western Australia Australian Journal of

Entomology 43(1) 88-93

Lodge DM 1993 Biological Invasions Lessons for ecology Trees 8 133-137

Louda SM and Collinge SK 1992 Plant resistance to insect herbivores A field test

of the environmental stress hypothesis Ecology 73 153-169

Lowman MD 1984 An assessment of techniques for measuring herbivory is

rainforest defoliation more intense than we thought Biotropica 16 264-268

Loxton I and Forster S 2000 Brigalow Research Station Technical Report 1999-

2000 Rep No Q100098 Queensland Beef Industry Institute Department of

Primary Industries Queensland Theodore

Lűckhoff HA 1964 Diseases of exotic plantation trees in the Republic of South Africa

FAOIUFRO Symposium Meet VI

Lundquist JE and Purnell RC 1987 Effects of Mycosphaerella leaf spot on growth

of Eucalyptus nitens Plant Disease 71 1025-1029

Luque J Girbal J 1989 Dieback of cork oak (Quercus suber) in Catalonia (NE Spain)

caused by Botryosphaeria stevensii European Journal of Forest Pathology

19(1) 7ndash13

Luque J Parlade J and Pera J 2002 Seasonal changes in the susceptibility of

Quercus suber to Botryosphaeria stevensii and Phytophthora cinnamomi Plant


Macauley BJ and Fox LR 1980 Variation in total phenols and condensed tannins in

Eucalyptus leaf phenology and insect grazing Austral Ecology 5(1) 31-35

Madeira MV Fabiatildeo A Pereira JS Arauacutejo MC and Ribeiro C 2002 Changes in

carbon stocks in Eucalyptus globulus Labill plantations induced by different

water and nutrient availability Forest Ecology and Management 171(1-2) 75-85

286

Majer JD Reecher HF Wellington AB Woinarski JCZ and Yen AL 1997

Invertebrates of eucalypt formations In lsquoEucalypt Ecology Individuals to

Ecosystemsrsquo (eds E Williams and JCZ Woinarski) pp 278-302 Cambridge

University Press Cambridge

Manion EG and Zhang S 1989 Eucalyptus dunnii potential in the Peoplersquos Republic

of China In lsquoProceedings Fourth Technical Exchange Seminar China-Australia

Afforestationrsquo pp 20-24 Project at Dongmen State Forest Farm

Manion PD 1981 Tree disease concepts Prentice-Hall Inc Englewood Cliffs NJ

399 p

Marco MA and Lopez JA 1995 Performance of Eucalyptus grandis and Eucalyptus

dunnii in the Mesopotamia region Argentina In lsquoEucalyptus Plantations

Improving Fibre Yield and Qualityrsquo (eds BM Potts NMG Boralho JB Reid RN

Cromer WN Tibbits CA Raymond) pp 40-45 Proceedings CRCTHF ndash IURFO

Conference Hobart CRC for Temperate Hardwood Forestry Hobart

Marks GC Fuhrer BA and Walters NEM 1982 In lsquoTree Disease in Victoriarsquo

Forest Commission Victoria Handbook No 1rsquo (Forests commission Melbourne)

Marks GC and Minko G 1969 The pathogenicity of Diplodia Pinea on Pinus radiata

d Don Australian Journal of Botany 17 1-12

Matheson AC and Cotterill PP 1990 Utility of genotype x environment interactions

Forest Ecology and Management 30 159-174

Matthews EG and Reid CAM 2002 A guide to the genera of the beetles of South

Australia In lsquoPart 8 Chrysomelidaersquo pp 66 South Australian Museum Adelaide

Mattson W Hack R 1987 In lsquoThe role of drought stress in provoking outbreaks of

phytophagous insectsrsquo Academic Press London

Mauchline N Withers T M Wang Q and Davis L1999 Life history and abundance

of the Eucalyptus leafroller Strepsicrates macropetana Meyrick pp 108-112

Proc 52nd New Zealand Plant protection Conference

Maxwell DL Kruger EL and Stanosz GR 1997 Effects of water stress on

Colonization of Poplar stems and excised leaf disks by Septoria musiva


287

Maxwell FG and Jennings PR 1980 Breeding Plants Resistant to Insects pp 683

John Wiley and Sons New York

Maxwell A Dell B Neumeister-Kemp HG and Hardy GEStJ 2003

Mycosphaerella species associated with Eucalyptus in south-western Australia

new species new records and a key Mycological Research 107(3) 351-359

Mazanec Z 1974 Influence of jarrah leaf miner on the growth of jarrah Australian

Forestry 37 32-42

McInnes RS and Carne PB 1978 Predation of Cossid Moth Larvae by Yellow-

Tailed Black Cockatoos Causing Losses in Plantations of Eucalyptus Grandis in

North Coastal New South Wales Australian Wildlife Research 5(1) 101 ndash 121

McClure MS 1980 Foliar nitrogen a basis for host suitability for elongate hemlock

scale Fiornia externa Ecology 61 72-79

McDonald GI 1981 Differential defoliation of Douglas fir trees by western spruce

budworm USDA Forestry Service Intemin Note INT-30610

McGrath JF 1999 Silviculture management options for E globulus plantations In

lsquoBalancing Productivity and Drought in Blue Gum Plantationsrsquo Proceedings f a

workshop presented by Bunnings Tree Far Department of Conservation and

Land Management CSIRO Forestry and Forest Products and Timber Eucalypts

Ltd Pemberton Western Australia 9-10 November 1999 (eds S Crombie J

McGrath and DA White) pp 23-27 Department of Conservation and Land

Management Perth

McPartland JM 1983 Stress Predisposition and Histopathology of Canker Diseases

in Woody Hosts MS Thesis University Illinois Urbana pp 60

McQuillan PB 1985 A taxonomic revision of the autumn gum moth genus

Mnesampela Guest (Lepidoptera Geometridae Ennominae) Entomology of

Scandinavia 16 175-202

Mendel Z Protasov A Blumberg D Saphir N Madar Z and La Salle J 2007

Release and recovery of the parasitoids of the eucalypt gall wasp Ophelimus

maskelli in Israel Phytoparasitica 35(4) 330-332

Metaliaj R Sicoli G and Luisi N 2003 Pathogenicity of Armillaria spp on water-

stressed Mediterranean oak seedlings Journal of Plant Pathology 85(4) 311

288

Milgate AW Potts BM Joyce H Mohammed C and Vaillancourt RE 2005

Genetic variation in Eucalyptus globulus for susceptibility to Mycosphaerella

nubilosa and its association with tree growth Australasian Plant Pathology 34

11-18

Milgate AW Yuan ZQ Vaillancourt R E and Mohammed C 2001

Mycosphaerella species occurring on Eucalyptus globulus and Eucalyptus nitens

plantations in Tasmania Australia Forest Pathology 31 53-63

Miles PW Aspinall D and Correl AT 1982 The response of two chewing insects on

water stressed food plants in relation to changes in their chemical composition

Australian Journal of Zoology 30 347-355

Miller DR and Wallner WE 1989 Influence of Climate on Gypsy Moth Defoliation In

Southern New England Environmental Entomology 18(4) 646-650

Mitchell CE and Power AG 2003 Release of invasive plants from fungal and viral

pathogens Nature 421 625-627

Mohammed C Wardlaw T Smith S Pinkard E Battaglia M Glen M Tommerup

I Potts B and Vaillancourt R 2003 Mycosphaerella leaf diseases of temperate

eucalypts around the southern Pacific Rim New Zealand Journal of Forestry

Science 33 362-372

Monteith GB 1991a The life and times of the giant wood moth Wildlife Australia

28(1) 8-10

Monteith GB 1991b lsquoLook whorsquos emerging ndash the birth of a giant wood moth Wildlife

Australia 28(2) 19

Moore LM and Wilson LF 1983 Recent advances in research of some pest

problems of hybrid Populus in Michigan and Wisconsin United States

Department of Agriculture and Forestry Services Technical Report NC-91

Moricca S 2002 Phomopsis alnea the cause of dieback of black alder in Italy Plant


Morrow PA 1977 The significance of phytophagous insects in the Eucalyptus forests

of Australia In lsquoThe Role of Arthropods in Forest Ecosystemsrsquo (eds WJ

Mattson) pp 19-29 Springer-Verlag New York

289

Morrow PA and Fox L R 1980 Effects of variation of eucalyptus essential oil yield

on insect growth and grazing damage Oecologia 45 209-219

Morrow PA Whitham TG Potts PM Ladiges P Ashton DH and Williams JB

1994 Gall forming insects concentrate on hybrid phenotypes of eucalyptus In

rsquoThe Ecology and Evolution of Gall forming Insectsrsquo (eds PW Price WJ

Mattson YN Baranchikov) pp 121-34 Forest Service General Technical

Report NC 174 United States Department of Agriculture St Paul MN

Munsell Albert H (1905) A Color Notation (ed G H Ellis) Boston USA

Nag Raj T R 1993 Coelomycetes anamorphs with appendage bearing conidia In

lsquoMycologue Publicationsrsquo Waterloo Canada

Nahrung HF Dunstan PK and Allen GR 2001 Larval gregariousness and neonate

establishment of the eucalypt-feeding beetle Chrysophtharta agricola

(Coleoptera Chrysomelidae Paropsini) Oikos 94 358-364

Nahrung HF 2006 Paropsine beetles (Coleoptera Chrysomelidae) in South-East

Queensland hardwood plantations identifying potential pest species Australian

Forestry 69 270-274

Nair KSS 2001 Pest outbreaks in tropical forest plantations Is there a greater risk for

exotic tree species Centre for international forestry research Indonesia

National Forestry Inventory (2007) National Plantation Update - March 2007 Bureau of

Rural Sciences Canberra

New T 1943 Evolution origins and Importance of insect-plant associations In lsquo

Associations between insects and plantsrsquo pp 1-14 (NSW University Press)

Nichol NS Wingfield MJ and Swart WJ 1992a Differences in susceptibility of

Eucalyptus species to Phaeoseptoria eucalypti European Journal of Forest


Nichol NS Wingfield MJ and Swart WJ 1992b The effect of site preparation and

and fertilisation on the severity of Phaeoseptoria eucalypti on eucalypt species

European Journal of Forest Pathology 22 424-431

Nielsen ES Edwards ED and Rangsi TV 1996 In lsquoChecklist of the Lepidoptera of

Australiarsquo CSIRO Melbourne

290

Nikles DG Lee DJ Robson K J Ponroy PC and Walker SM 2000 Progress

on species selection trials and genetic improvement of hardwoods for

commercial plantings in Queensland In lsquoAFG 2000 conference Opportunities for

the new Millenniumrdquo (eds A Snell and S Vize) pp 33-31 Australian Forest

Growers Cairns Queensland

Nixon KM and Hagedorn SF 1984 A Eucalyptus species and provenance trail on

two sites in the Natal Midlands Wattle Research Institute Report for 1983-1984

Thirty Seventh year September pp 134-137

Noble IR 1989 Ecological traits of the Eucalyptus LrsquoHerit Subgenera Monocalyptus

and Symphyomyrtus Australian Journal of Botany 37 207-224

Nuttall MJ 1983 Strepsicrates macropetana Meyrick (Lepidoptera Tortricidae)

Eucalyptus leafroller New Zealand Forest Service Forest and Timber Insects in

New Zealand No 57

Nylander JAA Ronquist F Huelsenbeck JP and Nieves-Aldrey JL 2004

Bayesian Phylogenetic Analysis of Combined Data Systematic Biology 53(1)

47-57

Obrycki JJ and Kring TJ 1998 Predacious Coccinellidae in biological control

Annual Review of Entomology 43 295-321

Ohmart CP Stewart LG and Thomas RJ 1983a Phytophagous insects

communities in the canopies of three Eucalyptus forest types in south east

Australia Australian Journal of Ecology 8 395-403

Ohmart CP Stewart LG and Thomas RJ 1983b Leaf consumption by insects in

three Eucalyptus forest types in Southeastern Australia and their role in short

term nutrient cycling Oecologia 59 322-330

Ohmart CP Thomas RJ and Stewart LG 1985 Effects of food quality particularly

nitrogen concentrations of Eucalyptus blakelyi foliage on the growth of Paropsis

atomaria larvae (Coleoptera Chrysomelidae) Oecologia 65(4) 543-549

Ohmart CP Thomas RJ and Stewart LG 1987 Nitrogen leaf toughness and the

population dynamics of Paropsis atomaria Oliver (Coleoptera Chrysomelidae) A

Hypothesis Journal of the Australian Entomological Society 26 203-207

291

Ohmart CP and Edwards PB 1991 Insect herbivory on Eucalyptus Annual Review

of Entomology 36 637-657

Old K M 1990 Diseases caused by fungi In lsquoTrees for Rural Australiarsquo (ed KW

Cremer) pp 210-216 Inkuta Press Melbourne

Old KM and Davison EM 2000 Canker diseases of eucalypts In lsquoDiseases and

Pathogens of Eucalyptsrsquo (Eds PJ Keane GA Kile FD Podger BN Brown) pp

241-258 CSIRO Publishing Melbourne

Old KM Gibbs R Craig I Myers BJ and Yuan QZ 1990 The effect of drought

and defoliation on the susceptibility of eucalypts to cankers caused by Endothia

gyrosa and Botryosphaeria ribis Australian Journal of Botany 38 571-581

Old K M Murray DIL Kile JA Simpson J and Malafant KWJ 1986 The

pathology of fungi isolated from eucalypt cankers in south-east Australia Journal

of Australian Forestry Research 16 21-36

Old KM Wingfield MJ and Yuan ZQ 2003 lsquoA Manual of Diseases of Eucalypts in

South-East Asiarsquo ACIAR Canberra and CIFOR Bogor

Old KM Yuan QZ and Kobayashi T 1991 A Valsa teleomorph of Cytospora

eucalypticola Mycological Research 95 1253-1256

Oliveira JG 1988 Eucalyptus tree improvement program at Rigesa In JG Carneiro et

al (ed) Bilateral symposium Brazil- Finland on Forestry actualities Curitiba

Parana Brazil

Orshan G 1954 Surface reduction and its significance as a hydrological factor Journal

of Ecology 42 442-444

Ostry ME and McNabb HSJr 1983 Diseases of intensely cultivated hybrid poplars

A summary of recent research in the north central region United States


Ostry ME and McNabb HSJr 1986 Poplus species and hybrid clones resistant to

Melampsora Marssonina and Septoria United States Department of Agriculture

and Forestry Services Technical Report NC-272

Paine TD Raffa KF Harrington TC 1997 Interactions among scolytid bark

beetles their associated fungi and live host conifers Annual Review of


292

Painter RH 1951 In lsquoInsect Resistance in Crop Plantsrsquo pp 520 University Press

Kansas Lawrence and London

Park RF and Keane PJ 1982a Three Mycosphaerella species from leaf diseases of

Eucalyptus Transactions of the British Mycological Society 79(1) 95-100

Park RF and Keane PJ 1982b Leaf diseases of Eucalypts associated with

Mycosphaerella species Transactions of the British Mycological Society 79(1)

101-115

Park RF Keane PJ Wingfield MJ and Crous PW 2000 Fungal disease of

eucalypt foliage In lsquoDiseases and Pathogens of Eucalyptsrsquo (eds PJ Keane GA

Kile FD Podger BN Brown) pp 153-239 CSIRO Publishing Melbourne

Patel JD 1971 Morphology of the gum tree scale Eriococcus coriaceus Maskell

(Homoptera Eriococcidae) with notes on it life history and habits near Adelaide

South Australia Australian Journal of Entomology 10(1) 43ndash56

Paton DM 1981 Eucalyptus Physiology III Frost Resistance Australian Journal of

Botany 29 675-88

Paulin-Mahady AE Harrington TC and McNew D 2002 Phylogenetic and

taxonomic evaluation of Chalara Chalaropsis and Thielaviopsis anamorphs

associated with Ceratocystis Mycologia 94 62-72

Pearce MH Malajczuk N1990 Factors affecting growth of Armillaria luteobubalina

rhizomorphs in soil Mycological Research 94(1) 38-48

Pegg G Brown B and Ivory M 2003 Eucalypt diseases in hardwood plantations in

Queensland Report no 16 Hardwoods Queensland Forestry Research

Department of Primary Industries Queensland Government

Pegg G Carnegie AJ Drenthe A and Wingfield MJ 2005 Quambalaria pitereka on

spotted gum plantations in Queensland and northern New South Wales

Australia The International Forestry Review 7(5) 337

Pegg G OrsquoDwyer C Carnegie AJ Burgess TI Wingfield MJ and Drenth A

2008 Quambalaria species associated with eucalypt plantation development


293

Pereira JCD Higa AR Shimivu JY and Higa RCV 1986 Comparison of the

wood provenances of Eucalyptus dunnii for energy purposes Boletim de

Perquisa Florestal 13 9-16

Philpott A 1923 Spilonata macropetana in New Zealand New Zealand Journal of

Science and Technology 6 216-217

Philips C 1992a Eucalyptus Weevil PIRSA Forestry No 7

Philips C 1992b Leafhoppers PIRSA Forestry No 2

Phillips CL 1993 Insect pest problems of eucalypt plantations in Australia 5 South


Pook EW Gill AM and Moore PHR 1998 Insect herbivory in a Eucalyptus

maculata forest on the south coast of New South Wales Australian Journal of

Botany 46 735-742

Protasova A Blumberga D Brandb D La Sallec J and Mendel Z 2007 Biological

control of the eucalyptus gall wasp Ophelimus maskelli (Ashmead) Taxonomy

and biology of the parasitoid species Closterocerus chamaeleon (Girault) with

information on its establishment in Israel Biological Control 42(2) 196-206

Price PW 1991 The plant vigour hypothesis and herbivore attack Oikos 62 244-51

Punithalingham E and Waterson JM 1970 Diplodia Pinea CMI Descriptions of plant

pathogenic fungi and Bacteria No173 Commonwealth Mycological Institute

Association of Applied Biology Key Surrey England

Purnell RC and Lundquist JE 1986 Provenance variation in Eucalyptus nitens on

the eastern Transvaal highveld in South Africa South African Forestry Journal

138 23-31

Rand TA 1999 Effects of environmental context on the susceptibility of Atriplex patula

to attack by herbivorous beetles Oecologia 121 39-46

Rao MR Singh MP and Day R 2001 Insect pest problems in tropical agroforestry

systems Contributory factors and strategies for management Journal

Agroforestry Systems 50(3) 243-277

Rausher MD 1981 The effect of native vegetation on the susceptibility of Aristolochia

reticulata (Aristolochiacea) to herbivore attack Ecology 62 1187-1195

294

Rayner ADM Boddy L 1988 Fungal Decomposition of Wood Its Biology and

Ecology Chichester UK New York USA Brisbane Australia Toronto Canada

Singapore Malaysia John Wylie and Sons

Rentz DCF 1996 The Abundant Orthopteroid Insects of Australia In lsquoGrasshopper

Countryrsquo University of New South Wales Press Australia

Richardson KF and Meakins RH 1986 Inter- and Intra-specific variation in the

susceptibility of eucalypts to the snout beetles Gonipterus scuttelatus Gyll

(Coleoptera Curculionidae) South African Journal of Forestry 139 21-31

Riek EF 1970 Chapter 29 Mantodea In lsquoThe Insects of Australiarsquo (ed DF

Waterhouse) pp 472-492 Melbourne University Press Melbourne Australia

Risch SJ Andow D and Alteiri MA 1983 Agroecosystem diversity and pest control

Data tentative conclusions and new research directions Environmental


Rivera AC Carbone SS and Andreacutes JA 2001 Life cycle and biological control of

the Eucalyptus snout beetle (Coleoptera Curculionidae) by Anaphes nitens

(Hymenoptera Mymaridae) in north-west Spain Agricultural and Forest


Roane MK Stipes RJ Phillips PM and Miller OKJr 1974 Endothia gyrosa

casual pathogen of pin oak blight Mycologia 66 1042- 1047

Ronquist F and Huelsenbeck JP 2003 MrBayes 3 Bayesian phylogenetic inference

under mixed models Bioinformatics 19(12) 1572-1574

Root RB 1973 Organisation of a plant-Arthropod association in simple and diverse

habitats The fauna of Collards (Brassicae oleraceae) Ecological Monographs

43(1) 95-124

Rosenheim JA Limburg DD and Colfer RG 1999 Impact of Generalist Predators

on a biological control agent Chrysoperla carnea Direct Observations

Ecological Applications 9(2) 409-417

Ross EW 1964 Cankers Associated with Ash Dieback Phytopathology 54 272-275

Ross EW 1966 Ash Dieback Etiological and developmental studies New York State

College of Forestry Technical Publication 88 80

295

Roux J Mthalana BL de Beer ZW and Wingfield MJ 2006 Quambalaria leaf and

shoot blight on Eucalyptus in South Africa Australasian Plant Pathology 35

427ndash33

Rozario SA Farrow RA and Gullan PJ 1993 Effects of ant attendance on

reproduction and survival of Euremeloides punctata (Signoret) and Eurymela

distincta (Signoret) (Hemiptera Eurymelidae) on eucalypts Australian Journal of

Entomology 32(2) 177ndash186

Rubtsov V V and Utkina I A (2003) Interrelations of green oak leaf roller population

and common oak Results of 30-year monitoring and mathematical modelling

Ecology Survey and Management of Forest Insects Proceeding 311 90-97

Sands PJ Rwlins W and Battaglia M 1999 Use of a simple plantation productivity

model to study the profitability of irrigated Eucalyptus globulus Ecological

Modelling 117 125-141

Sankaran KV Sutton BC and Minter DW 1995 A checklist of fungi recorded on

eucalypts Mycological Papers 170 1-376

Sanson G and Read J Aranwela N Clissold F Peeters P 2001 Measurement of

leaf biochemical properties in studies of herbivory Opportunities problems and

procedures Austral Ecology 26 535-546

Sauvard D 2000 In lsquoBark and Wood Boring Insects in Living Trees in Europe a

Synthesisrsquo (eds Lieutier F KR Day A Battisti JC Greacutegoire and H Evans) pp

63-68 Kluwer Academic Publishers Dordrecht The Netherlands

Schimel D S House1 J I Hibbard KA Bousquet P Ciais P Peylin PBH

Braswell MJ Apps D Baker A Bondeau J Canadell G Churkina1 W

Cramer AS Denning CB Field P Friedlingstein C Goodale M Heimann

RA Houghton JM Melillo B Moore D Murdiyarso I Noble SW Pacala

IC Prentice1 MR Raupach PJ Rayner RJ Scholes WL and Wirth SC

2001 Recent patterns and mechanisms of carbon exchange by terrestrial

ecosystems Nature 414 169-172

Schoener TW 1974 Resource Partitioning in Ecological Communities Science 185

27-39

296

Schoeneweiss DF 1975 Predisposition stress and plant disease Annual Review of


Schoeneweiss DF 1981 The role of environmental stress in diseases of woody

plants Plant Disease 65 308-314

Schowalter TD Hargrove WW and Crossley DA Jr 1986 Herbivory in forest

ecosystems Annual Review of Entomology 31 177-196

Schwerdtfeger F 1929 Ein Beitrag zur Fortpflanzungsbiologie des Borkenkafers

Pityogenes chalcographus L Z Angew Entomology 15 335-427

Schwerdtfeger F 1971 Vergleichende Untersuchungen an der Kronenfauna der

Eichen in Latenz- und Gradationsgebieten des Eichenwicklers (Tortrix viridana

L) Zeitschrift fbull ur angewandte Entomologie 67 296-304

Self NM Aitken EAB and Dale MD 2002 Susceptibility of provenances of spotted

gums to ramularia shoot blight New Zealand Plant Protection 55 68ndash72

Selman BJ1994 The evolutionary biology and taxonomy of Australian eucalyptus

beetles Entomography 3 451-454

Sharma PJ and Crowden RK 1974 Anthocyanins in some Eucalyptus species

Australian Journal of Botany 22 623-627

Shear CL Stevens NE and Tiller RJ 1917 Endothia parasitica and related

species United States Department of Agriculture Bulletin 380 1-82

Shearer BL 1994 The major plant pathogens occurring in natural ecosystems of

south-western Australia Journal of the Royal Society of Western Australia 77

113-122

Shearer BL Tippett JT and Bartle JR 1987 Botryosphaeria ribis infection

associated with death of Eucalyptus radiata in species selection trials Plant

Disease 71 140-145

Shivas RG 1989 Fungal and bacterial diseases of plants in Western Australia

Journal of the Royal Society of Western Australia 72 1-62

Simpson JA 2000 Quambalaria a new genus of eucalypt pathogens Australasian

Mycologist 19 57-62

297

Simmul TL and de Little DW 1999 Biology of the Paropsini (Chrysomelidae

Chrysomelinae) In lsquoAdvances in Chrysomelidae Biologyrsquo (ed Cox ML) pp 463-

477 Blackhuys Publishers Leiden

Sivanesan A and Shivas RG 2002 Studies on Mycosphaerella species in

Queensland Australia Mycological Research 106 355-364

Slatyer RO 1967 In lsquoPlant Water Relationshipsrsquo pp 366 New York Academic Press

New York

Slippers B Fourie G Crous PW Coutinho TA Wingfield BD Carnegie AJ and

Wingfield MJ 2004 Speculation and distribution of Botryosphaeria spp on

native and introduced Eucalyptus trees in Australia and South Africa Studies in

Mycology 50 343-358

Smith H Kemp G H J and Wingfield M J 1994 Canker and die-back of Eucalyptus

in South Africa caused by Botryosphaeria dothidea Plant Pathology 43(6)

1031ndash1034

Southcott RV 1978 Lepidopterism in the Australian region Records of the Adelaid

Childrens Hospital 2 67-73

South East Queensland Drought to 2007 2007 Department of Natural Resources and

Water Queensland

Soria F and Borralho NMG 1997 The genetics of resistance to Phoracantha

semipunctata attack in Eucalyptus globulus in Spain Silvae Genetica 46(6)

365-369

Specht RL Specht A Whelan MB and Hegarty EE 1995 In lsquoConservation Atlas

of Plant Communities in Australiarsquo Southern Cross University Press Lismore

Speight R and Wylie F 2001 In lsquoInsect Pests of Tropical Forestryrsquo pp 370 Cabi

New York

Steinbauer MJ and Clarke M 1998 Oviposition preference of a Eucalyptus herbivore

and the importance of leaf age on interspecific host choice Ecological


Steinbauer MJ 2001 The ecology research and management implications of insect

eucalypt interactions Symposium introduction Austral Ecology 26 445-446

298

Stipes RJ and Phillips PM 1971 A species of Endothia associated with a canker

disease of pin oak (Quercus palustris) in Virginia Plant Disease 55 467-469

Stolzy LH Letey J Klotz LJ and Labanauskas CK 1965 Water and aeration as

factors in root decay of Citrus sinensus Phytopathology 55 270-275

Stone C 1993 Fertilizer and insecticide effects on tree growth and psyllid infestation of

young Eucalyptus grandis and E dunnii plantations in northern New South

Wales Australian Forestry 33(1) 51ndash56

Stone C 1991 Insect attack of eucalypt plantations and regrowth forests in New South

Wales ndash A discussion paper Forest Commission of New South Wales Forest

Resource Series No 17

Stone C and Bacon PE 1994a Insect Herbivory in a River Red Gum (Eucalyptus

camaldulensis Dehnh) Forest in Southern New South Wales Australian Journal


Stone C and Bacon PE 1994b Relationships amongst moisture stress insect

herbivory foliar cineole content and the growth of river red gum Eucalyptus

camaldulensis Journal of Applied Ecology 31 604-612

Stone C and Bacon PE 1995 Leaf dynamics and insect herbivory in Eucalyptus

camaldulensis forest under moisture stress Journal of Applied Ecology 20 473-

81

Stone C and Birk E 2001 Benefits of weed control and fertiliser application to young

Eucalyptus dunnii stressed from water logging and insect damage Australian

Forestry Journal 64 151-158

Stone C 2001 Reducing the impact of insect herbivory in eucalypt plantations through

management of intrinsic influences on tree vigour Austral Ecology 26 482-488

Stone C Matsuki M and Carnegie AJ 2003 Pest and disease assessment in young

eucalypt plantations Field manual for using the crown damage index (ed M

Parsons) Natural Forest Inventory Bureau of Rural Sciences Canberra

Australia

Stone JK and Petrini O 1997 Endophytes of forest trees a model for fungus-plant

interactions In lsquoThe Mycota V Plant Relationships Part Brsquo (eds GC Carroll and

P Tudzynski) pp 129-142 Springer and Verlag

299

Stork NE 1988 Insect diversity Facts fiction and speculation Biological Journal of

the Linnean Society 35(4) 321-337

Strauss SY 2001 Benefits and risks of biotic exchange between Eucalyptus

plantations and native Australian forests Austral Ecology 26(5) 447-457

Strauss SY and Agrawal AA 1999 The ecology and evolution of plant tolerance to

herbivory Tree 14 179-185

Strozaker R Lefroy T Keating B and Williams J 2000 A revolution in land use

emerging land use systems for managing dryland salinity pp 24 CSIRO Land

and Water Canberra

Suleman P Al-Musallam A MenezesCA 2001 The effect of solute potential and

water stress on black scorch caused by Chalara paradoxa and Chalara radicicola

on date palms Plant Disease 1 80-83

Surico G Mugnai L Pastorelli R Giovannetti L Stead DE 1996 Erwinia alni a

new species causing bark cankers of alder (Alnus Miller) species International

Journal of Systematic Bacteriology 46 720ndash6

Sutton BC 1971 Coelomycetes IV The genus Harknessia and similar fungi on


Sutton BC 1975 Eucalyptus microfungi Satchmopsis gen nov and new species of

Coniella Coniothyrium and Harknessia Nova Hedwiga 26 1-16

Sutton BC and Pascoe IG 1989 Addenda to Harknessia (Coelomycetes)


Swart W I Knowx-Davies P S and Wingfield M J 1985 Sphaeropsis sapinea with

special reference to its occurrence on Pinus spp in South Africa South African


Swart WJ and Conradie E 1992 Effects of water stress on the development of

cambial lesions caused by Cryphonectria cubensis on Eucalyptus grandis Plant

Disease 76(7) 744-746

Swofford DL Waddell PJ Huelsenbeck PJ and Foster PG 2001 Bias in

phylogenetic estimation and its relevance to the choice between parsimony and

likelihood methods Systematic Biology 50 (4) 525-539

300

Tanton MT and Khan SM 1978 Aspects of the biology of the eucalypt-defoliating

chrysomelid beetle Paropsis atomaria OI in the Australian Capital Territory

Australian Journal of Zoology 26(1) 113 ndash 120

Taylor GS 1997 Effect of plant compounds on the population dynamics of the lerp

insect Cardiaspina albitextura Taylor (Psylloidea Spondyliaspididae) on

eucalypts In lsquoEcology and Evolution of Plant Feeding Insects in Natural and

Manmade Environmentsrsquo (ed A Raman) pp 37-57 International Scientific

Publications New Dehli

Thomson GE 1941 Leaf spot diseases of Poplars caused by Septoria musiva and S

populicola Phytopathology 31 241-254

Thomson VP Nicotra AB and Steinbauer MJ 2001 Influence of previous frost

damage on tree growth and insect herbivory of Eucalyptus globulus globulus


Thumlert TA and Austin AD 1994 Biology of Phylacteophaga froggati Riek

(Hymeoptera Pergidae) and its parasitoids in South Australia Transactions of the

Royal Society of South Australia 118 99-113

Tibbits WN 1986 Eucalypt plantations in Tasmania Australian Forestry 49 219-223

Tippett JY Crombie DS and Hill TC 1987 Effect of phloem water relations on the

growth of Phytophthora cinnamomi Phytopathology 77 246-250

Tippett YJ McGrath JF and Hill TC 1989 Site and seasonal effects on

susceptibility of Eucalyptus marginata to Phytophthora cinnamomi Australian

Journal of Botany 37 481-490

Timberlake PH 1957 A new Entodontine Chalcid-fly from seed capsules of

Eucalyptus in Califormia (Hymenoptera Eulophidae) Pan-Pacific Entomologist

33 109-110

Tinsley TW 1953 The effects of varying the water supply to plants on their

susceptibility to infection with viruses Annual Applied Biology 40 750-760

Tobiessen P and Buchsbaum S 1976 Ash Dieback and Drought Canadian Journal of

Botany 54 543-545

Todd JW 1989 The ecology and behavious of Nezara viridula Annual Review of


301

Took FGC 1955 The eucalyptus snout beetles Gonipterus scuttelatus Gylenhall A

study of its control by biological means Entomological Memoirs 3 1-281

Turnbull JW 2000 Economic and social importance of eucalypts In lsquoDiseases and

Pathogens of Eucalyptsrsquo (eds PJ Keane GA Kile FD Podger BN Brown)

pp 1-9 CSIRO Publishing Melbourne

Urquhart CA and Stone C 1995 In lsquoPsyllids in Eucalypt Plantationsrsquo Forest

Protection Research Division Series No E3 State Forests of New South Wales

Beecroft NSW

Valentini VA 1994 Influence of water relations on Quercus cerris-Hypoxylon

mediterraneum interaction a model of drought-induced susceptibility to a

weakness parasite Tree Physiology 14(2) 129-139

Van der Kamp BJ 1991 Pathogens as agents of diversity in tropical landscapes

Forestry Chronicle 67 353-354

van Heerden SW and Wingfield MJ 2002 Effect of environment on the response of

Eucalyptus clones to inoculation by Cryphonectria cubensis Forest Pathology

32 295-402

Venter M Wingfield MJ Countinho TA and Wingfield BB 2001 Molecular

characterisation of Endothia gyrosa isolates from Eucalyptus in South Africa and

Australia Plant Pathology 50 211-217

Venter M Myburg H Wingfield BD Coutinho TA and Wingfield MJ 2002 A

new species of Cryphonectria from South Africa and Australia pathogenic to

Eucalyptus Sydowia 54 98-117

Vinaya Rai RS Parthiban KT and Kumaravelu G 1995 Studies on the drought

tolerance of Eucalyptus at seedling stage Journal of Tropical Forest Science

8(2) 155-160

Volker PW Owen JV and Borralho NMG 1994 Genetic variances and

covariences for frost tolerance in Eucalyptus globulus and E nitens Silviculture

Genetics 43 366-372

Vranjic JA and Gullan PJ 1990 The Effect of a Sap-Sucking Herbivore Eriococcus

coriaceus (Homoptera Eriococcidae) on Seedling Growth and Architecture in

Eucalyptus blakelyi Oikos 59(2) 157-162

302

Wainwright M Swan HT 1986 CG Paine and the earliest surviving clinical records

of penicillin therapy Medical History 30(1) 42ndash56

Waldboth M Oberhuber W 2009 Synergistic effect of drought and chestnut blight

(Cryphonectria parasitica) on growth decline of European chestnut (Castanea

sativa) Forest Pathology 39(1) 43ndash55

Walker J 1962 Notes on plant parasitic fungi I Proceedings of the Linnean Society of

New South Wales 87 162-176

Walker J and Bertus AL 1971 Shoot blight of Eucalyptus spp caused by an

undescribed species of Ramularia Proceedings of the Linnean Society of New

South Wales 96 108-115

Walker J Old KM and Murray DIL 1985 Endothia gyrosa on Eucalyptus in

Australia with notes on other species of Endothia and Cryphonectria Mycotaxon

23 350-370

Walker J Sutton BC and Pascoe IG 1992 Phaeoseptoria eucalypti and similar

fungi on Eucalyptus with description of Kirramyces gen nov (Coelomycetes)


Walker JC and Stahmann MA 1955 Chemical nature of disease resistance Annual

Review of Plant Physiology 6 351-366

Walklate PJ McCartney HA and Fitt BDL 1989 Vertical dispersal of plant

pathogens by splashing Part II experimental study of the relationship between

raindrop size and the maximum splash height Plant Pathology 38(1) 64-70

Wang HR and Zhou WL 1996 Fertiliser and eucalypt plantations in China In

lsquoNutrition of Eucalyptsrsquo (eds PM Attiwill and MA Adams MA) pp 389-397

CSIRO Melbourne

Wardlaw TJ 1999 Endothia gyrosa associated with severe stem cankers on

plantations grown Eucalyptus nitens in Tasmania Journal of Forest Pathology

29 199-208

Wargo PM 1996 Consequences of environmental stress on oak predisposition to

pathogens Annals of Forest Science 53 (2-3) 359-368

303

Waring GL and Cobb NS 1992 The impact of plant stress on herbivore population

dynamics In lsquoInsect Plant Interactionsrsquo Vol 4 (ed E Bernays) pp 167-226 CRC

Press Boca Roton

Waring GL and Price PW 1988 Consequences of host plant chemical and physical

variability to an associated herbivore Ecological Research 3 205-216

Waterhouse DF 1970 In lsquoThe Insects of Australiarsquo A Textbook for Students and

Research Workers and the Supplement First Edition Melbourne University

Press

Waterson D 1995 Gumleaf Skeletoniser Forest Protection Research Division Series

No E7 State Forests of New South Wales Beecroft NSW

Waterson D and Urquhart CA 1995 Leaf beetles Forest Protection Research

Division Series No E6 State Forests of New South Wales Beecroft NSW

Wermelinger B 2004 Ecology and management of the Spruce Bark Beetle Ips

typhographus a review of recent research Forest Ecology and Management

202 67-82

Weston CJ Attiwill PM and Cameron JN 1991 Growth and nutrition of eucalypts

in relation to soil type and former land use in Gippsland Victoria In lsquoIntensive

Foresty The Role of Eucalyptsrsquo IUFRO symposim (eds APG Schonau) pp

480-491 South African insititute of Forestry Durban

White DA 1996 In lsquoPhysiological responses to drought of Eucalyptus globulus and E

nitens in plantationsrsquo PhD Thesis University of Tasmania 168 pp

White DA and Kile GA 1993 Discolouration and decay from artificial wounds in 20

year old Eucalyptus regnans European Journal of Forest Pathology 23 431-

440

White T Bruns T Lee S and Taylor J 1990 Amplification and direct sequencing of

fungal ribosomal RNA genes for phylogenetics In lsquoPCR protocols a Guide to

Methods and Applicationsrsquo (Eds M Innis D Gelfand J Snisky and T White) pp

315-322 (Academic Press San Diego)

White TCR 1969 An index to measure weather induced stress of trees associated

with outbreaks of psyllids in Australia Ecology 50 905-9

304

White TCR 1974 A hypothesis to explain outbreaks of looper caterpillars with special

reference to populations of Selidosema suavis in a plantation of Pinus radiata in

New Zealand Oecologia 16 279-301

White TCR 1984 The abundance of invertebrate herbivores in relation to the

availability of nitrogen in stressed food plants Oecologia 63 90-105

White TCR 1969 An index to measure weather-induced stress of trees associated


White TCR 1986 Weather Eucalyptus dieback in New England and a general

hypothesis of the cause of dieback Pacific Science 40 58-78

Whitham TG 1989 Plant hybrid zones as sinks for insect pests Science 244 1490-

1493

Whitham TG Morrow PA and Potts BM 1994 Plant hybrid zones as centres for

biodiversity The herbivore community of two endemic Tasmanian eucalypts

Oecologia 97 481-490

Whyte G 2002 Insect-Fungal Relationships on Eucalyptus camaldulensis in the

Gresswell Forest Reserve Bundoora Melbourne Honours Thesis La Trobe

University

Whyte G Burgess TI Barber PA and Hardy GESt J 2005 First record of

Mycosphaerella heimii in Australia Australasian Plant Pathology 34 605-606

Wingfield M J Crous PW and Boden K 1996 Kirramyces destructans sp nov a

serious leaf pathogen of Eucalyptus in Indonesia South African Journal of

Botany 62 325-327

Wingfield MJ Crous PW and Couthinho TA 1997 A serious new canker disease

of Eucalyptus in South Africa caused by a new species of Coniothyrium

Mycopathologia 136 139-146

Wingfield MJ 2001 Worldwide movement of exotic forest fungi especialy in the

tropics and the southern hemisphere Bioscience 51 134-139

Winjum JK Dixon RK and Schroeder PE 1993 Forest management and carbon

storage An analysis of 12 key forest nations Water Air and Pollution 70(1-4)

239-257

305

Withers TM 2001 Colonization of eucalypts in New Zealand by Australian insects


Withers TM Raman A and Berry JA 2000 Host range and biology of Ophelmius

eucalypti (Gahan) (Hymenoptera Eulophidae) A pest of New Zealand Eucalypts

New Zealand Plant Protection 53 339-344

Wood DL 1982 The role of pheromones kairomones and allomones in the host

selection and colonization behaviour of bark beetles Annual Review of


Woodward TE Evans JW and Eastop VF 1970 Chapter 26 Hemiptera In lsquoThe

Insects of Australiarsquo (Ed DF Waterhouse) pp 387-457 (Melbourne University

Press)

Wylie FR Johnsston PJM and Eismann RL 1993 A survey of native tree dieback

in Queensland Research Paper no 16 Department of Primary Industries

Queensland

Wylie FR and Peters BC 1993 Insect pest problems of eucalypt plantations in

Australia Queensland Australian Forestry 56 358-362

Xu D and Dell B 1997 Importance of micronutrients for productivity of plantation

eucalypts in east Asia In lsquoProceedings of 6th Annual BIO-REFOR Workshoprsquo lsquo(ed

J Kikkawa) pp 133-138 Brisbane Queensland BIO-REFOR University of

Tokyo Tokyo

Xu D Dell B Malajczuk N and Gonga M 2002 Effects of P fertilisation on

productivity and nutrient accumulation in a Eucalyptus grandis times E urophylla

plantation in southern China Forest Ecology and Management 161 89-100

Yamamura K and Kiritani K 1998 A simple method to estimate the potential increase

in the number of generations under global warming in temperate zones Applied

Entomological Zoology 33 289-298

Yarwood CE 1959 Predisposition In lsquoPlant Pathologyrsquo (eds JG Horsfall AE

Diamond) pp 674 New York and London Academic New York

Yuan ZQ 1989 Mycology and pathology of seed-borne fungi of Australian native

trees and of eucalypt canker fungi Msc Thesis Xinjiang Agricultural University

Urumqui P R China

306

Yuan ZQ 1998 Stem canker diseases of eucalypts in Tasmania PhD Thesis

University of Tasmania Hobart Australia

Yuan ZQ 1999 In lsquoFungi Associated with Diseases Detected during Health Surveys of

Eucalypt Plantations in Tasmaniarsquo PhD Thesis School of Agricultural Science

University of Tasmania Hobart

Yuan ZQ and Mohammed C 1997a Investigation of fungi associated with stem

cankers of eucalypts in Tasmania Australia Australian Plant Pathology 26 78-

84

Yuan ZQ and Mohammed C 1999 Pathogenicity of stem cankers associated with

Eucalyptus in Tasmania Australia Plant Disease 83 1063-1069

Yuan ZQ and Mohammed C 2000 The pathogenicity of isolates of Endothia gyrosa

to Eucalyptus nitens and E globulus Australasian Plant Pathology 29 29-35

Yuan ZQ and Mohammed C 2001 Lesion development in stems of rough and

smooth barked Eucalyptus nitens following artificial inoculations with canker

fungi Forest Pathology 31 149-161

Zalucki MP Anthony RC and Malcolm BS 2002 Ecology and behaviour of first

instar larval Lepidoptera Annual Review of Entomology 47 361-393

Zangerl AR Arntz AM and Berenbaum MR 1997 Physiological price of an

induced chemical defence photosynthesis respiration biosynthesis and growth


Zanuncio TV Zanuncio JC Miranda MMM and Medeiros AGD 1998 Effect of

plantation age on diversity and population fluctuation of Lepidoptera collected in

Eucalyptus plantations in Brazil Forest Ecology and Management 108 91-98

Zhang L Dowling T Hocking M Morris J Adams G Hickel K Best A and

Vertessy R 2003 Predicting the effects of large-scale afforestation on annual

flow regime and water allocation an example for the Goulburn-Broken

catchments Technical report 035 Cooperative Research Centre for Catchment

Hydrology

Zhonghua M Morgan DP and Michailides TJ 2001 Effect of water stress on

Botryosphaeria blight of pistachio caused by Botryosphaeria dothidia Plant

Disease 85 745-749

ii

Declaration

I declare that this thesis is my own account of research and contains as its main content

work that has not previously been submitted for a degree at any tertiary education

institution

Gilbert Whyte

April 2012

iii

Acknowledgments









my final years




encouragement


iv

INDEX









Defining Stress 8





Thesis Chapters 24



Introduction 26


Results 32


Eucalypt Weevils 37


Giant Wood Moths 45

Case Moths 48

Cup Moths 50

Leaf Bag Worms 53




Mirid Bugs 63

v



Psyllids 70

Leafhoppers 73

Planthoppers 75

Clown Bugs 77

Assassin Bugs 79

Ladybird Beetles 81

Praying Mantids 83

Lacewings 85

Discussion 87



Introduction 91


Results 101










Discussion 130



SEASON 134

Introduction 134


vi

Results 153

Discussion 184



Introduction 191


Results 198

Discussion 216



Introduction 221


Results 231

Discussion 237


Important Pests 242







Future Research 257


8 REFERENCES 259

vii

Abstract
























viii


























ix











1























2


Inventory 2007)























3














4

























5
























6











Grafton

Bundaberg

Brisbane

Rockhampton

Coffs Harbour

Urbenville




QLD-NSW

Border


7






















8



Defining Stress





















9

























10

























11

























12





al 2006)
















groups (Table 11)

13


Borers






Sauvard 2004


Abies spp

Larix spp

Pinus spp

Picea spp













Christiansen 1992



14

Borers continued



Eucalyptus spp

Acacia spp





Duffy 1963

Hanks et al 1999




Pinus banksiana

Pinus strobes

Picea abies





Lavallee 1994


Larix decidua

Picea spp

Pinus spp




Schwerdtfeger 1929

Avtzis et al 2000

Kirisits 2004


Pinus sp

Picea sp

Europe

North West Africa

Northern Asia







15

Defoliating Insects






Miles et al 1982


Quercus spp

Tsuga canadensis

Europe

Asia

North America





Davidson et al 1999


Cedrus spp

Larix spp

Pinus spp




Rouault et al 2006

Buffo et al 2007


Quercus spp

Acer spp

Betula spp

Fagus spp

Populus spp





Gasow 1925

Schwerdtfeger 1971

Larsson et al 2000


16

Other Insects





White 1969

Miles et al 1982

17

























18
























19










Wargo 1996

Metaliaj 2003


Fagus silvatica

and Quercus spp



Hendry et al 1998









Barr 1972


Zhonghua et al 2001


20



Phoenix dactylifera

Saudi Arabia

Iraq




Suleman et al 2001



Quercus spp

Castanopsis spp

Acer spp

Rhus spp

Typhina spp

Carya ovata

Europe

Asia

Africa

North America





Shear et al 1917

Hepting 1974

Anagnostakis1984




Ross 1964




Britain



Gibbs et al 1997


Acer spp

Populus spp




Christensen 1940

Bertrand 1967

Jones 1985

Guyon et al1996

21



Australia




Shearer et al 1987

Old et al 1990


Quercus spp

Juniperus spp

Fraxinus spp

Eucalyptus spp

Europe

South America

North America




Luque et al 2002


Pinus spp

Picea spp

Abies spp




Birch 1937

Laughton1937

Eldridge 1961

Lűckhoff 1964

Buchanan 1967



Barker 1979

Gibson 1980

Brown et al 1981

Chou 1982

Swart et al 1985

22



North America

Australia

South Africa




Schoenweiss 1975

Davison 1982


Walker et al 1985

Roane et al 1986

Old et al 1990

Gryzenhout 2006



Quercus spp

Castanea spp

Populus spp






Valentini 1994

Neofusicoccum ribis


E dunnii

E grandis

E camaldulensis

E radiata

E cladocalyx

E marginata

Corymbia calophylla




Shearer et al 1987

Old et al 1990

Luque et al 2002

Slippers et al 2004

23



Southeast Asia

Africa



Surico et al1996

Moricca 2002


Aspen spp

North America





Bier 1939

Thomson 1941




Abebe and Hart 1990

Maxwell 1997

Xylella fastidiosa


and Citrus spp



Boyer 1995

24





development


southern Queensland


Queensland region



plantation age)





pathogens



Thesis Chapters




25






Southern Queensland


Southern Queensland






Plantations


26


Introduction











Stone 2001)











27


























28


























29




1985 Farr 2002)











development

Chapter Aim





presented

30


Site Selection




South Wales






grandis)

Sampling Regime











31

















32

Results










P

P


Oxyops sp

Defoliator

Defoliator

Medium Low

P

P

P

P

P

P

P

P


Paropsis obsoleta

Paropsis variolosa



Paropsisterna sp

Cryptocephalus sp

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

High

Low

Low

High

Low

Low

Low

Low



P

P


Defoliator

Defoliator

Low

Low










P

P


Cardiaspina sp

Sap-sucker

Sap-sucker

Medium

Low



P

P


Amorbus sp

Sap-sucker

Sap-sucker

Low

Medium








33














34

Longicorn Beetles

Order Coleoptera

Family Cerambycidae















2-1B amp C)







(Lawson et al 2002)

35












eventually die

Threat to Industry






Lawson et al 2002)






36





37

Eucalypt Weevils

Order Coleoptera
























38










Threat to Industry













39


40

Chrysomelid Beetles

Order Coleoptera
























41




Biology and Ecology











Symptoms and Damage




Threat to Industry






42















43


44


45

Giant Wood Moths

Order Lepidoptera

Family Cossidae























46

















Threat to Industry






47


48

Case Moths

Order Lepidoptera

Family Xyloryctidae





















49

Threat to Industry








50

Cup Moths

Order Lepidoptera

Family Limacodidae









Common 1990)














51





Symptoms and Damage



Threat









minor pests

52


53

Leaf Bag Worms

Order Lepidoptera

Family Psychidae






















54















Threat to Industry










55




56


Order Lepidoptera

Family Tortricidae





















57






Threat to Industry










Queensland

58


59

Leaf Blister Sawfly

Order Hymenoptera



















1984)





60


Threat to Industry








61

Eucalypt Gall Wasps

Order Hymenoptera























62


Threat to Industry









Queensland


63

Mirid Bugs

Order Hemiptera

Family Miridae

Species Rayieria sp












mimics

Symptoms and Damage




Threat to Industry




64








65

Brown Scale Insects

Order Hemiptera

Family Eriococcidae















1971)







66






Threat to Industry




Wales









67


68


Order Hemiptera

Family Pentatomidae






















69

Threat to Industry




references







closely


70

Psyllids

Order Hemiptera

Family Psyllidae























71


1974)

Threat to Industry















72


73

Leafhoppers

Order Hemiptera

Family Eurymelidae








visible dorsally












of insects occur

74

Threat to Industry










75

Planthoppers

Order Hemiptera

Family Flatidae

Species Siphanta sp















Threat to Industry





76


77

Clown Bugs

Order Hemiptera

Family Coreidae


















2-18B)




78

Threat to Industry





1998)






79

Assassin Bugs

Order Hemiptera

Family Reduviidae
















Role in Plantations






80


81

Ladybird Beetles

Order Coleoptera











(Figure 2-20B)







Role in Plantations




82


83

Praying Mantids

Order Mantodea










Rentz 1966)







Role in Plantations




84


85

Lacewings

Order Neuroptera










D) (Riek 1970)













86

Role in Plantations






87

Discussion























88





















Urquhart 1995)





89

























90






















91


Introduction






















92





Pegg et al 2005)


















(Cheah 1977 Fry 1983)



93














2003)












94


























95



et al 2004)

Chapter Aim






Site Selection







Sampling Regime





Sampling Method


96















Fungal Isolation










97













dark at 20C












98














1990)











99
















Phylogenetics









100






















101

Results










Figure 3-12)


102


Saprophytic fungi




Aspergillus sp

Cladosporium sp

Epicoccum sp

Fusarium sp

Mucor sp

Penicillium sp

Pestalotiopsis sp


Phoma glomerata

Phomopsis diaporthe

Nigrasporum sp

Trichoderma sp












103






Primary Pathogens
















104


Primary Pathogens










105



Field Symptoms




(Figure 3-2A amp B)

Growth on Agar









Ecology and threat







106






plantation industry

107


108


Host E dunnii

Field Symptoms





Growth on Agar











spore (Figure 3-3G)

Ecology and Threat




109








110


A

B

111


Host E globulus

Field Symptoms





Growth on Agar










Ecology and Threat






112




113


Host E grandis

Field Symptoms





Growth on Agar









Ecology and Threat






114



115


Host E globulus

Field Symptoms





Growth on Agar










Ecology and Threat






116




117


Host E dunnii

Field Symptoms




ostioles

Growth on Agar











Ecology and Threat



Queensland

118


119

New Fungal Species















(Table 32)

120







MUCC467 EU300999

MUCC468 EU301000

MUCC649 DQ240133

MUCC693 EU301002

MUCC694 DQ240169


























MUCC668 EU301011

MUCC669 EU301014

































5 changes

100

67

100

100

76

100

75

96

100

55

100

100

100

99

92

85

87

57

100100

99

100

100

100

100

97

100

100

86

98

66

97

91

97

100

84

84

99

98

88

52





121



















122

























123








124


125

















3-11 E)




ex-type MUCC649)




126






127













(Figure 3-12 K)









MUCC648)



128




129


130

Discussion







species

















Asci

131


















132













conditions






glasshouse






133















Conclusion







134


Introduction





















135

























136

























137

















Chapter Aim








138




Site Selection





















139


Size (ha)














E

S

N

W




140

Group - Age

Size (ha)








Fertiliser History

















141

















defined (Table 43)

142



Foliar Yellowing



Foliar Reddening



143





Total Fungal Damage



144









145





Mirid Damage



146


Psyllid Damage






147





Weevil Defoliation




148


Foliar Wasp Galls



Scale Insect Damage



149





150











of each plantation






151








the average leaf






152







plantation groups
















153


















Results

Damage Averages






154



recorded










Mirid Damage 29 9th







Total 100










155

Plantation Age


34 60 43 53

Local Climate


37 58

Seasons


41 30 59 60















South-4 01 0660 01 0231 01 0179








156













North-3



South-1

South-2

South-3

South-4

North-1

North-2

North-4

North-3

157










158



1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th

13th











1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th

13th

14th

15th











159







Table 412)

160



mean 23 18 1 34 14

plusmn SE 14 1 07 44 19

mean 3 3 13 19 16 16 01

plusmn SE 13 13 08 37 13 13 02

mean 19 16 08 28 04 04 01

plusmnSE 16 11 07 45 09 09 02

mean 19 16 08 28 04 04 01

plusmnSE 08 08 1 46 02

mean 22 2 12 27 05 05 04

plusmnSE 13 12 09 42 1 1 11 01

mean 19 16 08 28 04 04


mean 184 183 116 28 28

plusmn SE 6 59 58 41 41

mean 121 12 121 125 04 04

plusmn SE 57 57 67 354 06 06

mean 145 13 25 13 03

plusmn SE 29 21 46 35 05

mean 116 111 116 37 03 08 08 01


Plantation mean 152 138 152 43 06 06 13 09 24

Group plusmn SE 79 79 66 37 12 12 23 2 4

mean 519 222 519 33 02 09 3 03

plusmn SE 74 74 68 7 05 27 69 05

mean 191 143 191 53 01 01 16 04

plusmnSE 116 68 116 45 02 02 26 07

mean 421 155 421 56 02

plusmnSE 75 75 53 72 04

mean 321 165 321 46 02 02 04 09 07 06 02

plusmnSE 173 78 173 56 06 06 12 21 35 22 04

mean 152 152 175 313 22 05 05

plusmnSE 45 45 32 17 87 06 06

mean 371 371 124 7 02 02 88 13

plusmn SE 72 72 25 35 04 04 12 35

mean 174 174 9 71 05 05

plusmn SE 58 58 17 14 07 06

mean 245 245 113 41 77

plusmn SE 85 85 41 82 83

mean 235 235 126 88 15 03 03 22 03

plusmnSE 18 18 43 16 96 05 05 69 18


Nov-04

1

2

1

2

3

4

4

Total

1


Total

3

4

Total

1

2

3

2

3

4

Total

AgeEstate month

May-05

Feb-05

Aug-04


161



mean 33 13 26 38 378 01

plusmn SE 28 28 25 16 165 04

mean 16 154 141 163 32 179 177 145 19 01

plusmn SE 52 48 46 85 17 5 5 46 37 04

mean 135 13 155 222 141 93 92 146 16 01 05

plusmn SE 68 68 52 179 15 4 4 37 35 02 08

mean 343 341 145 43 94 67 65 38 39

plusmn SE 64 64 45 137 18 48 48 5 4

mean 168 159 117 26 229 85 83 83 18 01

plusmnSE 13 13 67 183 185 76 75 75 34 02 05

mean 9 9 55

plusmn SE 55 55 09

mean 9 9 15 44 24 23 147

plusmn SE 38 38 72 73 24 24 69

mean 29 29 75 25 3 3 02 13

plusmn SE 51 51 1 46 22 22 07 11

mean 172 171 92 29 67 67 01

plusmn SE 22 22 44 76 39 39 02

Northern mean 139 139 93 24 29 29 38 03


Group mean 238 238 5 06 58 58 11 163 06

plusmn SE 25 2 18 72 72 18 132 18

mean 35 35 15 63 78 77 26 01

plusmn SE 113 113 34 92 57 55 4 02

mean 34 34 111 17 17 16 14 02 01

plusmn SE 125 125 43 29 09 09 24 04 02

mean 356 356 8 13 05 05 03

plusmn SE 57 57 2 23 07 07 07

mean 321 321 86 25 39 39 03 47 02 04 01

plusmnSE 99 98 37 53 53 53 1 95 09 13 04 02

mean 17 17 193 16 09 03 145 47 38 12 03

plusmn SE 43 43 56 31 27 07 8 14 58 2 09

mean 443 443 12 3 04 04 15 04

plusmn SE 33 33 2 3 09 05 11 04

mean 36 36 16 2 16 01 13 03

plusmn SE 62 62 17 28 35 03 19 09

mean 464 464 13 16 03 02

plusmn SE 87 87 27 23 04 04

mean 359 359 133 21 07 03 36 41 09 04 02

plusmnSE 131 131 49 27 22 05 02 74 83 32 11 05


Total

Total

2

Nov-04

1

2

1

2

3

4

3

4

1

2

3

4

Total

1

3

4

Total


month AgeEstate

May-05

Aug-04

Feb-05


162




















163



















164

Plantations Estates


1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th





Climate Averages







(Figure 4-6C amp D)









165








0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

Mean Rainfall (mm)

Mean Max Temp (C)

Mean Min Temp (C)


Months Months


0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300M

ean

Rain

fall (

mm

)

Mean Rainfall (mm)

Mean Max Temp (C)

Mean Min Temp (C)

START OF SURVEY

C D

2004 2005 2005 2004

A B

166










P R P R P R P R

Aug 04

Nov 04 01 0438

Feb 05 01 0631 01 0547

May 05 01 091 01 0934 01 077













167







Four Groupings

Stress 017

168

Rank

Seasons


1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th










169















plantation group




170


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










171


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










172



















categories






173


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










174


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










175
















176



00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D


G H










177


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H











178





Damage category







179


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










180




00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










181

Mirid Damage












182


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










183
























less than 1)

184

Discussion

























185








and season


















186













infestations













187









group











trees






188





















Humphreys 1965)





189














dieback












190






Economic Impacts









191


Introduction












and Wingfield 1996)











192



Queensland)
























193








(Dungey et al 2000)

Chapter Aim
















194









195


E dunnii





E grandis

(Flooded gum)




E globulus

(Blue gum)




E tereticornis

(Forest red gum)






196


E camaldulensis

(River red gum)




E urophylla





Hybrid taxa













197



Sampling Regime


Climate Data



and Figure 4-6)








198











those differences

Results









199













Total 100

Comparing Taxa















damage



th) caused by


200





201






Pure Taxa


1st

2nd

3

rd

4th

5th





Hybrid Taxa


1st

2nd

3

rd

4th

5th

6th








202















G1 G2

G3

G4

203










2004



P R P R P R P R

Aug 04

Nov 04 01 0763

Feb 05 01 0634 01 0271

May 05 01 0757 01 0481 01 0562





(Table 56)

Rank

Seasons of Sampling


1st

2nd

3

rd

4th

5th







204
















205












206

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)



A B C D

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)



A B C D

207






Total Fungal Damage








208

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)



A B C D

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u


Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)



A B C D

209

Foliar Yellowing












210



00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt TaxaP

erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)A B C D



00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

211





212

Aug-04

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 92 196 97 326 264 313 292 149

plusmn SE 07 14 17 34 00 23 47 21


M 92 196 94 326 250 313 292 149

plusmn SE 60 60 42 42 42 42 42 42


M 132 120 88 163 183 219 198 167

plusmn SE 17 13 19 24 23 18 21 23


M 09 172 14 00 00 00 00 00

plusmn SE 21 201 24 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Fungal Damage

M 144 150 94 288 83 167 410 128

plusmn SE 17 11 28 45 29 15 37 19

Foliar Yellowing

M 174 00 139 56 42 00 56 111

plusmn SE 63 00 77 56 42 00 56 77

Scale Insect Damage

M 00 00 03 00 00 00 00 00

plusmn SE 00 00 05 00 00 00 00 00


213

Nov-04

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 264 135 205 178 104 188 354 128

plusmn SE 19 12 21 29 07 09 51 08


M 264 135 25 177 14 188 354 128

plusmn SE 60 60 42 42 42 42 42 42


M 214 384 125 104 83 125 208 87

plusmn SE 15 25 00 14 14 00 51 07


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 51 00 00 00 00 00 00

plusmn SE 00 100 00 00 00 00 00 00

Total Fungal Damage

M 00 121 42 21 00 00 111 00

plusmn SE 00 34 23 07 00 00 51 00

Foliar Yellowing

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


214

Feb-05

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 373 167 146 417 125 458 436 413

plusmn SE 22 12 07 34 05 24 25 30


M 373 161 146 417 125 458 431 413

plusmn SE 60 60 42 42 42 42 42 42


M 175 239 146 153 125 167 156 125

plusmn SE 08 20 07 13 00 15 17 00


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 00 03 00 00 00 00 00

plusmn SE 00 00 06 00 00 00 00 00

Total Fungal Damage

M 30 00 10 340 00 292 188 264

plusmn SE 08 00 06 43 00 15 37 47

Foliar Yellowing

M 36 00 28 00 00 00 14 56

plusmn SE 29 00 19 00 00 00 14 33

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


215

May-05

E dunnii

E globulus

E grandis

E tereticornis






M 307 416 340 00 07 63 10 00

plusmn SE 66 65 85 00 05 16 11 00

Total Defoliation

M 429 117 219 244 66 417 549 444

plusmn SE 22 12 37 21 04 15 27 15


M 429 116 219 243 66 417 549 444

plusmn SE 60 60 42 42 42 42 42 42


M 98 70 101 69 63 63 267 163

plusmn SE 08 03 15 05 00 00 38 27


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 111

plusmn SE 00 00 00 00 00 00 00 192


M 02 00 00 00 00 00 00 00

plusmn SE 04 00 00 00 00 00 00 00

Total Fungal Damage

M 73 193 49 03 00 00 10 10

plusmn SE 21 23 27 04 00 00 06 06

Foliar Yellowing

M 00 00 00 14 00 333 83 42

plusmn SE 00 00 00 14 00 123 58 23

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


216

Discussion
























217












McGrath 1999)














218



Comparing Taxa



















oviposition of eggs



219









Conclusion

















220














221


Introduction






















222


























223























and Mohammed 1999)



224





















Chapter Aim




225






expression





tissue



226
























227




(White et al 1990)






Site Selection















228


















229



















230


1B Control
















experiments

Measuring Lesions











231







Statistics




treatment

Results


either produced a






232















W

L

233

64)











1A Control 20

2A H eucalypti 40

3A N ribis 40








00

1000

2000

3000

4000

5000

6000

Control

H eucalypti

B ribis

C eucalyptic

ola

H eucalypti +

B ribis

H eucalypti +

C e

ucalypticola

B ribis + C

euca

lyptic

ola

H eucalypt +

B ribis

+ C e

ucalypticola

Treatments

Mean

Lesio

n S

everi

ty I

nd

ex (

len

gth

x w

idth

) m

m LSD (5) = 12210


1A

2A

3A

4A

5A

6A 7A

8A

234











1B Control 20

2B 1 H eucalypti 90


4B 3 H eucalypti 80

5B 1 N ribis 95

6B 2 N ribis 80

7B 3 N ribis 50

8B 4 N ribis 70

9B 5 N ribis 70






235























0

50

100

150

200

250

300

350

400

450

500

Con

trol

1 H

euca

lypt

i

2 H

euca

lypt

i

3 H

euca

lypt

i

1 B

rib

is

2 B

rib

is

3 B

rib

is

4 B

rib

is

5 B

rib

is

1 C

euca

lypt

icol

a

2 C

euca

lypt

icol

a

3 C

euca

lypt

icol

a

4 C

euca

lypt

icol

a

Isolate species

Lesio

n S

everi

ty I

nd

ex (

len

gth

x w

idth

)

mm

LSD (5) = 7500

1B

2B 3B

4B

5B

6B

7B

8B 9B

10B 11B

12B 13B

236














treatments

Fungal Species


00

1000

2000

3000

4000

5000

6000

Cont

rol

H e

ucalyp

ti

B r

ibis

C e

ucalyp

ticola

Fungal species

Mean

Lesio

n S

everi

ty I

nd

ex (

len

gth

x

wid

th)

mm


Neofusicoccum ribis


Winter

Summer

237








treatment




Discussion













238


























239

























240


rainfall










Conclusion













241



242


Important Pests






















243

Important Pathogens
















al 2005)







high rainfall)

244

Economic Impacts

























245







future
















drought


246










al 2002)















247

inciting factors











248








illustrated





249








250




251















Threshhold

Pivot Threshhold

Pivot

Threshhold Pivot

252


















Threshhold Pivot

Threshhold Pivot

Threshhold Pivot

253




as follows




















254



deficiency



1 Drought Impacts


















255
























256







4 Sampling Regime










5 Weather Data







257

more efficient

Future Research















and adults






258

2 Leaf Pathogens






3 Canker Pathogens








Concluding Remarks






259

8 References






74






85-110








1-44








260









13 197-205






















261






167














346






155-163





106

262













Research 49 153-174













49 239-42





263






1318-1320





101 1060-1064





















264






173





370














Canberra






Japan

265











Ottawa



Entomology 9 65-72













Washington DC





266



44




Management 54 69-87























267







21 (2) 53-63
























268


London















Sciences 12 438-441















269



10 537-586


Plymouth UK






















15 409-416


no 74109rsquo

270






97



278



Paul Minnesota USA












441-458





532



271








HMSO London


10-11



















272



Science 63 597-612


















Bulletin 4 41-94











273



Workshop





Science 199 941-945







95 551-557


Mycology 41 339-339





Wales Beecroft NSW









274




of Ecology 12(2) 125-138










Dilemma FAO Rome








Zoology 28 83-90




23(4) 497-507



Science 21 185-193

275







25





Research 94 313-319








33 (2) 111ndash114









276




Publishing











Forestry 28 394-411












Forestry 91 215-221





277


Management 26 38-40





series No 4













Mycology 55 35-52







35-52





278




10-11





The Netherlands













153-162









279



353ndash361






565


1 31-50


67





Extension Nevada


Press Inc Hong Kong


Melbourne









280





















53







ACT



281












119










258 CSIRO Australia










282




















Netherlands





York





283



129-147




















277-83







284















No 982727 pp 28-38


London Academic














285




165-185



















19(1) 7ndash13









286









399 p






















287







Forestry 37 32-42














Management Perth











288




11-18














Science 33 362-372


28(1) 8-10


Australia 28(2) 19









289
















Forestry 69 270-274















290













New Zealand No 57



47-57















291




















Parana Brazil












292







101-115








Botany 29 675-88















293












Botany 46 735-742











138 23-31








294
























43(1) 95-124







295



427ndash33



























27-39

296






















113-122



Disease 71 140-145




Mycologist 19 57-62

297







New York




Mycology 50 343-358



1031ndash1034




Water Queensland



365-369




New York






298



















81









Australia




299









and Water Canberra


















Disease 76(7) 744-746




300

























33 109-110




Botany 54 543-545



301








Beecroft NSW








32 295-402









8(2) 155-160



Genetics 43 366-372




302













23 350-370











CSIRO Melbourne



29 199-208



303



Press Boca Roton





Press







202 67-82









440







304











1493






University





Botany 62 325-327








239-257

305











Press)



Queensland






Tokyo Tokyo











Urumqui P R China

306








84




















Hydrology



Disease 85 745-749

iii

Acknowledgments









my final years




encouragement


iv

INDEX









Defining Stress 8





Thesis Chapters 24



Introduction 26


Results 32


Eucalypt Weevils 37


Giant Wood Moths 45

Case Moths 48

Cup Moths 50

Leaf Bag Worms 53




Mirid Bugs 63

v



Psyllids 70

Leafhoppers 73

Planthoppers 75

Clown Bugs 77

Assassin Bugs 79

Ladybird Beetles 81

Praying Mantids 83

Lacewings 85

Discussion 87



Introduction 91


Results 101










Discussion 130



SEASON 134

Introduction 134


vi

Results 153

Discussion 184



Introduction 191


Results 198

Discussion 216



Introduction 221


Results 231

Discussion 237


Important Pests 242







Future Research 257


8 REFERENCES 259

vii

Abstract
























viii


























ix











1























2


Inventory 2007)























3














4

























5
























6











Grafton

Bundaberg

Brisbane

Rockhampton

Coffs Harbour

Urbenville




QLD-NSW

Border


7






















8



Defining Stress





















9

























10

























11

























12





al 2006)
















groups (Table 11)

13


Borers






Sauvard 2004


Abies spp

Larix spp

Pinus spp

Picea spp













Christiansen 1992



14

Borers continued



Eucalyptus spp

Acacia spp





Duffy 1963

Hanks et al 1999




Pinus banksiana

Pinus strobes

Picea abies





Lavallee 1994


Larix decidua

Picea spp

Pinus spp




Schwerdtfeger 1929

Avtzis et al 2000

Kirisits 2004


Pinus sp

Picea sp

Europe

North West Africa

Northern Asia







15

Defoliating Insects






Miles et al 1982


Quercus spp

Tsuga canadensis

Europe

Asia

North America





Davidson et al 1999


Cedrus spp

Larix spp

Pinus spp




Rouault et al 2006

Buffo et al 2007


Quercus spp

Acer spp

Betula spp

Fagus spp

Populus spp





Gasow 1925

Schwerdtfeger 1971

Larsson et al 2000


16

Other Insects





White 1969

Miles et al 1982

17

























18
























19










Wargo 1996

Metaliaj 2003


Fagus silvatica

and Quercus spp



Hendry et al 1998









Barr 1972


Zhonghua et al 2001


20



Phoenix dactylifera

Saudi Arabia

Iraq




Suleman et al 2001



Quercus spp

Castanopsis spp

Acer spp

Rhus spp

Typhina spp

Carya ovata

Europe

Asia

Africa

North America





Shear et al 1917

Hepting 1974

Anagnostakis1984




Ross 1964




Britain



Gibbs et al 1997


Acer spp

Populus spp




Christensen 1940

Bertrand 1967

Jones 1985

Guyon et al1996

21



Australia




Shearer et al 1987

Old et al 1990


Quercus spp

Juniperus spp

Fraxinus spp

Eucalyptus spp

Europe

South America

North America




Luque et al 2002


Pinus spp

Picea spp

Abies spp




Birch 1937

Laughton1937

Eldridge 1961

Lűckhoff 1964

Buchanan 1967



Barker 1979

Gibson 1980

Brown et al 1981

Chou 1982

Swart et al 1985

22



North America

Australia

South Africa




Schoenweiss 1975

Davison 1982


Walker et al 1985

Roane et al 1986

Old et al 1990

Gryzenhout 2006



Quercus spp

Castanea spp

Populus spp






Valentini 1994

Neofusicoccum ribis


E dunnii

E grandis

E camaldulensis

E radiata

E cladocalyx

E marginata

Corymbia calophylla




Shearer et al 1987

Old et al 1990

Luque et al 2002

Slippers et al 2004

23



Southeast Asia

Africa



Surico et al1996

Moricca 2002


Aspen spp

North America





Bier 1939

Thomson 1941




Abebe and Hart 1990

Maxwell 1997

Xylella fastidiosa


and Citrus spp



Boyer 1995

24





development


southern Queensland


Queensland region



plantation age)





pathogens



Thesis Chapters




25






Southern Queensland


Southern Queensland






Plantations


26


Introduction











Stone 2001)











27


























28


























29




1985 Farr 2002)











development

Chapter Aim





presented

30


Site Selection




South Wales






grandis)

Sampling Regime











31

















32

Results










P

P


Oxyops sp

Defoliator

Defoliator

Medium Low

P

P

P

P

P

P

P

P


Paropsis obsoleta

Paropsis variolosa



Paropsisterna sp

Cryptocephalus sp

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

High

Low

Low

High

Low

Low

Low

Low



P

P


Defoliator

Defoliator

Low

Low










P

P


Cardiaspina sp

Sap-sucker

Sap-sucker

Medium

Low



P

P


Amorbus sp

Sap-sucker

Sap-sucker

Low

Medium








33














34

Longicorn Beetles

Order Coleoptera

Family Cerambycidae















2-1B amp C)







(Lawson et al 2002)

35












eventually die

Threat to Industry






Lawson et al 2002)






36





37

Eucalypt Weevils

Order Coleoptera
























38










Threat to Industry













39


40

Chrysomelid Beetles

Order Coleoptera
























41




Biology and Ecology











Symptoms and Damage




Threat to Industry






42















43


44


45

Giant Wood Moths

Order Lepidoptera

Family Cossidae























46

















Threat to Industry






47


48

Case Moths

Order Lepidoptera

Family Xyloryctidae





















49

Threat to Industry








50

Cup Moths

Order Lepidoptera

Family Limacodidae









Common 1990)














51





Symptoms and Damage



Threat









minor pests

52


53

Leaf Bag Worms

Order Lepidoptera

Family Psychidae






















54















Threat to Industry










55




56


Order Lepidoptera

Family Tortricidae





















57






Threat to Industry










Queensland

58


59

Leaf Blister Sawfly

Order Hymenoptera



















1984)





60


Threat to Industry








61

Eucalypt Gall Wasps

Order Hymenoptera























62


Threat to Industry









Queensland


63

Mirid Bugs

Order Hemiptera

Family Miridae

Species Rayieria sp












mimics

Symptoms and Damage




Threat to Industry




64








65

Brown Scale Insects

Order Hemiptera

Family Eriococcidae















1971)







66






Threat to Industry




Wales









67


68


Order Hemiptera

Family Pentatomidae






















69

Threat to Industry




references







closely


70

Psyllids

Order Hemiptera

Family Psyllidae























71


1974)

Threat to Industry















72


73

Leafhoppers

Order Hemiptera

Family Eurymelidae








visible dorsally












of insects occur

74

Threat to Industry










75

Planthoppers

Order Hemiptera

Family Flatidae

Species Siphanta sp















Threat to Industry





76


77

Clown Bugs

Order Hemiptera

Family Coreidae


















2-18B)




78

Threat to Industry





1998)






79

Assassin Bugs

Order Hemiptera

Family Reduviidae
















Role in Plantations






80


81

Ladybird Beetles

Order Coleoptera











(Figure 2-20B)







Role in Plantations




82


83

Praying Mantids

Order Mantodea










Rentz 1966)







Role in Plantations




84


85

Lacewings

Order Neuroptera










D) (Riek 1970)













86

Role in Plantations






87

Discussion























88





















Urquhart 1995)





89

























90






















91


Introduction






















92





Pegg et al 2005)


















(Cheah 1977 Fry 1983)



93














2003)












94


























95



et al 2004)

Chapter Aim






Site Selection







Sampling Regime





Sampling Method


96















Fungal Isolation










97













dark at 20C












98














1990)











99
















Phylogenetics









100






















101

Results










Figure 3-12)


102


Saprophytic fungi




Aspergillus sp

Cladosporium sp

Epicoccum sp

Fusarium sp

Mucor sp

Penicillium sp

Pestalotiopsis sp


Phoma glomerata

Phomopsis diaporthe

Nigrasporum sp

Trichoderma sp












103






Primary Pathogens
















104


Primary Pathogens










105



Field Symptoms




(Figure 3-2A amp B)

Growth on Agar









Ecology and threat







106






plantation industry

107


108


Host E dunnii

Field Symptoms





Growth on Agar











spore (Figure 3-3G)

Ecology and Threat




109








110


A

B

111


Host E globulus

Field Symptoms





Growth on Agar










Ecology and Threat






112




113


Host E grandis

Field Symptoms





Growth on Agar









Ecology and Threat






114



115


Host E globulus

Field Symptoms





Growth on Agar










Ecology and Threat






116




117


Host E dunnii

Field Symptoms




ostioles

Growth on Agar











Ecology and Threat



Queensland

118


119

New Fungal Species















(Table 32)

120







MUCC467 EU300999

MUCC468 EU301000

MUCC649 DQ240133

MUCC693 EU301002

MUCC694 DQ240169


























MUCC668 EU301011

MUCC669 EU301014

































5 changes

100

67

100

100

76

100

75

96

100

55

100

100

100

99

92

85

87

57

100100

99

100

100

100

100

97

100

100

86

98

66

97

91

97

100

84

84

99

98

88

52





121



















122

























123








124


125

















3-11 E)




ex-type MUCC649)




126






127













(Figure 3-12 K)









MUCC648)



128




129


130

Discussion







species

















Asci

131


















132













conditions






glasshouse






133















Conclusion







134


Introduction





















135

























136

























137

















Chapter Aim








138




Site Selection





















139


Size (ha)














E

S

N

W




140

Group - Age

Size (ha)








Fertiliser History

















141

















defined (Table 43)

142



Foliar Yellowing



Foliar Reddening



143





Total Fungal Damage



144









145





Mirid Damage



146


Psyllid Damage






147





Weevil Defoliation




148


Foliar Wasp Galls



Scale Insect Damage



149





150











of each plantation






151








the average leaf






152







plantation groups
















153


















Results

Damage Averages






154



recorded










Mirid Damage 29 9th







Total 100










155

Plantation Age


34 60 43 53

Local Climate


37 58

Seasons


41 30 59 60















South-4 01 0660 01 0231 01 0179








156













North-3



South-1

South-2

South-3

South-4

North-1

North-2

North-4

North-3

157










158



1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th

13th











1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th

13th

14th

15th











159







Table 412)

160



mean 23 18 1 34 14

plusmn SE 14 1 07 44 19

mean 3 3 13 19 16 16 01

plusmn SE 13 13 08 37 13 13 02

mean 19 16 08 28 04 04 01

plusmnSE 16 11 07 45 09 09 02

mean 19 16 08 28 04 04 01

plusmnSE 08 08 1 46 02

mean 22 2 12 27 05 05 04

plusmnSE 13 12 09 42 1 1 11 01

mean 19 16 08 28 04 04


mean 184 183 116 28 28

plusmn SE 6 59 58 41 41

mean 121 12 121 125 04 04

plusmn SE 57 57 67 354 06 06

mean 145 13 25 13 03

plusmn SE 29 21 46 35 05

mean 116 111 116 37 03 08 08 01


Plantation mean 152 138 152 43 06 06 13 09 24

Group plusmn SE 79 79 66 37 12 12 23 2 4

mean 519 222 519 33 02 09 3 03

plusmn SE 74 74 68 7 05 27 69 05

mean 191 143 191 53 01 01 16 04

plusmnSE 116 68 116 45 02 02 26 07

mean 421 155 421 56 02

plusmnSE 75 75 53 72 04

mean 321 165 321 46 02 02 04 09 07 06 02

plusmnSE 173 78 173 56 06 06 12 21 35 22 04

mean 152 152 175 313 22 05 05

plusmnSE 45 45 32 17 87 06 06

mean 371 371 124 7 02 02 88 13

plusmn SE 72 72 25 35 04 04 12 35

mean 174 174 9 71 05 05

plusmn SE 58 58 17 14 07 06

mean 245 245 113 41 77

plusmn SE 85 85 41 82 83

mean 235 235 126 88 15 03 03 22 03

plusmnSE 18 18 43 16 96 05 05 69 18


Nov-04

1

2

1

2

3

4

4

Total

1


Total

3

4

Total

1

2

3

2

3

4

Total

AgeEstate month

May-05

Feb-05

Aug-04


161



mean 33 13 26 38 378 01

plusmn SE 28 28 25 16 165 04

mean 16 154 141 163 32 179 177 145 19 01

plusmn SE 52 48 46 85 17 5 5 46 37 04

mean 135 13 155 222 141 93 92 146 16 01 05

plusmn SE 68 68 52 179 15 4 4 37 35 02 08

mean 343 341 145 43 94 67 65 38 39

plusmn SE 64 64 45 137 18 48 48 5 4

mean 168 159 117 26 229 85 83 83 18 01

plusmnSE 13 13 67 183 185 76 75 75 34 02 05

mean 9 9 55

plusmn SE 55 55 09

mean 9 9 15 44 24 23 147

plusmn SE 38 38 72 73 24 24 69

mean 29 29 75 25 3 3 02 13

plusmn SE 51 51 1 46 22 22 07 11

mean 172 171 92 29 67 67 01

plusmn SE 22 22 44 76 39 39 02

Northern mean 139 139 93 24 29 29 38 03


Group mean 238 238 5 06 58 58 11 163 06

plusmn SE 25 2 18 72 72 18 132 18

mean 35 35 15 63 78 77 26 01

plusmn SE 113 113 34 92 57 55 4 02

mean 34 34 111 17 17 16 14 02 01

plusmn SE 125 125 43 29 09 09 24 04 02

mean 356 356 8 13 05 05 03

plusmn SE 57 57 2 23 07 07 07

mean 321 321 86 25 39 39 03 47 02 04 01

plusmnSE 99 98 37 53 53 53 1 95 09 13 04 02

mean 17 17 193 16 09 03 145 47 38 12 03

plusmn SE 43 43 56 31 27 07 8 14 58 2 09

mean 443 443 12 3 04 04 15 04

plusmn SE 33 33 2 3 09 05 11 04

mean 36 36 16 2 16 01 13 03

plusmn SE 62 62 17 28 35 03 19 09

mean 464 464 13 16 03 02

plusmn SE 87 87 27 23 04 04

mean 359 359 133 21 07 03 36 41 09 04 02

plusmnSE 131 131 49 27 22 05 02 74 83 32 11 05


Total

Total

2

Nov-04

1

2

1

2

3

4

3

4

1

2

3

4

Total

1

3

4

Total


month AgeEstate

May-05

Aug-04

Feb-05


162




















163



















164

Plantations Estates


1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th





Climate Averages







(Figure 4-6C amp D)









165








0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

Mean Rainfall (mm)

Mean Max Temp (C)

Mean Min Temp (C)


Months Months


0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300M

ean

Rain

fall (

mm

)

Mean Rainfall (mm)

Mean Max Temp (C)

Mean Min Temp (C)

START OF SURVEY

C D

2004 2005 2005 2004

A B

166










P R P R P R P R

Aug 04

Nov 04 01 0438

Feb 05 01 0631 01 0547

May 05 01 091 01 0934 01 077













167







Four Groupings

Stress 017

168

Rank

Seasons


1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th










169















plantation group




170


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










171


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










172



















categories






173


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










174


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










175
















176



00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D


G H










177


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H











178





Damage category







179


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










180




00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










181

Mirid Damage












182


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










183
























less than 1)

184

Discussion

























185








and season


















186













infestations













187









group











trees






188





















Humphreys 1965)





189














dieback












190






Economic Impacts









191


Introduction












and Wingfield 1996)











192



Queensland)
























193








(Dungey et al 2000)

Chapter Aim
















194









195


E dunnii





E grandis

(Flooded gum)




E globulus

(Blue gum)




E tereticornis

(Forest red gum)






196


E camaldulensis

(River red gum)




E urophylla





Hybrid taxa













197



Sampling Regime


Climate Data



and Figure 4-6)








198











those differences

Results









199













Total 100

Comparing Taxa















damage



th) caused by


200





201






Pure Taxa


1st

2nd

3

rd

4th

5th





Hybrid Taxa


1st

2nd

3

rd

4th

5th

6th








202















G1 G2

G3

G4

203










2004



P R P R P R P R

Aug 04

Nov 04 01 0763

Feb 05 01 0634 01 0271

May 05 01 0757 01 0481 01 0562





(Table 56)

Rank

Seasons of Sampling


1st

2nd

3

rd

4th

5th







204
















205












206

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)



A B C D

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)



A B C D

207






Total Fungal Damage








208

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)



A B C D

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u


Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)



A B C D

209

Foliar Yellowing












210



00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt TaxaP

erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)A B C D



00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

211





212

Aug-04

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 92 196 97 326 264 313 292 149

plusmn SE 07 14 17 34 00 23 47 21


M 92 196 94 326 250 313 292 149

plusmn SE 60 60 42 42 42 42 42 42


M 132 120 88 163 183 219 198 167

plusmn SE 17 13 19 24 23 18 21 23


M 09 172 14 00 00 00 00 00

plusmn SE 21 201 24 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Fungal Damage

M 144 150 94 288 83 167 410 128

plusmn SE 17 11 28 45 29 15 37 19

Foliar Yellowing

M 174 00 139 56 42 00 56 111

plusmn SE 63 00 77 56 42 00 56 77

Scale Insect Damage

M 00 00 03 00 00 00 00 00

plusmn SE 00 00 05 00 00 00 00 00


213

Nov-04

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 264 135 205 178 104 188 354 128

plusmn SE 19 12 21 29 07 09 51 08


M 264 135 25 177 14 188 354 128

plusmn SE 60 60 42 42 42 42 42 42


M 214 384 125 104 83 125 208 87

plusmn SE 15 25 00 14 14 00 51 07


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 51 00 00 00 00 00 00

plusmn SE 00 100 00 00 00 00 00 00

Total Fungal Damage

M 00 121 42 21 00 00 111 00

plusmn SE 00 34 23 07 00 00 51 00

Foliar Yellowing

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


214

Feb-05

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 373 167 146 417 125 458 436 413

plusmn SE 22 12 07 34 05 24 25 30


M 373 161 146 417 125 458 431 413

plusmn SE 60 60 42 42 42 42 42 42


M 175 239 146 153 125 167 156 125

plusmn SE 08 20 07 13 00 15 17 00


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 00 03 00 00 00 00 00

plusmn SE 00 00 06 00 00 00 00 00

Total Fungal Damage

M 30 00 10 340 00 292 188 264

plusmn SE 08 00 06 43 00 15 37 47

Foliar Yellowing

M 36 00 28 00 00 00 14 56

plusmn SE 29 00 19 00 00 00 14 33

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


215

May-05

E dunnii

E globulus

E grandis

E tereticornis






M 307 416 340 00 07 63 10 00

plusmn SE 66 65 85 00 05 16 11 00

Total Defoliation

M 429 117 219 244 66 417 549 444

plusmn SE 22 12 37 21 04 15 27 15


M 429 116 219 243 66 417 549 444

plusmn SE 60 60 42 42 42 42 42 42


M 98 70 101 69 63 63 267 163

plusmn SE 08 03 15 05 00 00 38 27


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 111

plusmn SE 00 00 00 00 00 00 00 192


M 02 00 00 00 00 00 00 00

plusmn SE 04 00 00 00 00 00 00 00

Total Fungal Damage

M 73 193 49 03 00 00 10 10

plusmn SE 21 23 27 04 00 00 06 06

Foliar Yellowing

M 00 00 00 14 00 333 83 42

plusmn SE 00 00 00 14 00 123 58 23

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


216

Discussion
























217












McGrath 1999)














218



Comparing Taxa



















oviposition of eggs



219









Conclusion

















220














221


Introduction






















222


























223























and Mohammed 1999)



224





















Chapter Aim




225






expression





tissue



226
























227




(White et al 1990)






Site Selection















228


















229



















230


1B Control
















experiments

Measuring Lesions











231







Statistics




treatment

Results


either produced a






232















W

L

233

64)











1A Control 20

2A H eucalypti 40

3A N ribis 40








00

1000

2000

3000

4000

5000

6000

Control

H eucalypti

B ribis

C eucalyptic

ola

H eucalypti +

B ribis

H eucalypti +

C e

ucalypticola

B ribis + C

euca

lyptic

ola

H eucalypt +

B ribis

+ C e

ucalypticola

Treatments

Mean

Lesio

n S

everi

ty I

nd

ex (

len

gth

x w

idth

) m

m LSD (5) = 12210


1A

2A

3A

4A

5A

6A 7A

8A

234











1B Control 20

2B 1 H eucalypti 90


4B 3 H eucalypti 80

5B 1 N ribis 95

6B 2 N ribis 80

7B 3 N ribis 50

8B 4 N ribis 70

9B 5 N ribis 70






235























0

50

100

150

200

250

300

350

400

450

500

Con

trol

1 H

euca

lypt

i

2 H

euca

lypt

i

3 H

euca

lypt

i

1 B

rib

is

2 B

rib

is

3 B

rib

is

4 B

rib

is

5 B

rib

is

1 C

euca

lypt

icol

a

2 C

euca

lypt

icol

a

3 C

euca

lypt

icol

a

4 C

euca

lypt

icol

a

Isolate species

Lesio

n S

everi

ty I

nd

ex (

len

gth

x w

idth

)

mm

LSD (5) = 7500

1B

2B 3B

4B

5B

6B

7B

8B 9B

10B 11B

12B 13B

236














treatments

Fungal Species


00

1000

2000

3000

4000

5000

6000

Cont

rol

H e

ucalyp

ti

B r

ibis

C e

ucalyp

ticola

Fungal species

Mean

Lesio

n S

everi

ty I

nd

ex (

len

gth

x

wid

th)

mm


Neofusicoccum ribis


Winter

Summer

237








treatment




Discussion













238


























239

























240


rainfall










Conclusion













241



242


Important Pests






















243

Important Pathogens
















al 2005)







high rainfall)

244

Economic Impacts

























245







future
















drought


246










al 2002)















247

inciting factors











248








illustrated





249








250




251















Threshhold

Pivot Threshhold

Pivot

Threshhold Pivot

252


















Threshhold Pivot

Threshhold Pivot

Threshhold Pivot

253




as follows




















254



deficiency



1 Drought Impacts


















255
























256







4 Sampling Regime










5 Weather Data







257

more efficient

Future Research















and adults






258

2 Leaf Pathogens






3 Canker Pathogens








Concluding Remarks






259

8 References






74






85-110








1-44








260









13 197-205






















261






167














346






155-163





106

262













Research 49 153-174













49 239-42





263






1318-1320





101 1060-1064





















264






173





370














Canberra






Japan

265











Ottawa



Entomology 9 65-72













Washington DC





266



44




Management 54 69-87























267







21 (2) 53-63
























268


London















Sciences 12 438-441















269



10 537-586


Plymouth UK






















15 409-416


no 74109rsquo

270






97



278



Paul Minnesota USA












441-458





532



271








HMSO London


10-11



















272



Science 63 597-612


















Bulletin 4 41-94











273



Workshop





Science 199 941-945







95 551-557


Mycology 41 339-339





Wales Beecroft NSW









274




of Ecology 12(2) 125-138










Dilemma FAO Rome








Zoology 28 83-90




23(4) 497-507



Science 21 185-193

275







25





Research 94 313-319








33 (2) 111ndash114









276




Publishing











Forestry 28 394-411












Forestry 91 215-221





277


Management 26 38-40





series No 4













Mycology 55 35-52







35-52





278




10-11





The Netherlands













153-162









279



353ndash361






565


1 31-50


67





Extension Nevada


Press Inc Hong Kong


Melbourne









280





















53







ACT



281












119










258 CSIRO Australia










282




















Netherlands





York





283



129-147




















277-83







284















No 982727 pp 28-38


London Academic














285




165-185



















19(1) 7ndash13









286









399 p






















287







Forestry 37 32-42














Management Perth











288




11-18














Science 33 362-372


28(1) 8-10


Australia 28(2) 19









289
















Forestry 69 270-274















290













New Zealand No 57



47-57















291




















Parana Brazil












292







101-115








Botany 29 675-88















293












Botany 46 735-742











138 23-31








294
























43(1) 95-124







295



427ndash33



























27-39

296






















113-122



Disease 71 140-145




Mycologist 19 57-62

297







New York




Mycology 50 343-358



1031ndash1034




Water Queensland



365-369




New York






298



















81









Australia




299









and Water Canberra


















Disease 76(7) 744-746




300

























33 109-110




Botany 54 543-545



301








Beecroft NSW








32 295-402









8(2) 155-160



Genetics 43 366-372




302













23 350-370











CSIRO Melbourne



29 199-208



303



Press Boca Roton





Press







202 67-82









440







304











1493






University





Botany 62 325-327








239-257

305











Press)



Queensland






Tokyo Tokyo











Urumqui P R China

306








84




















Hydrology



Disease 85 745-749

iv

INDEX









Defining Stress 8





Thesis Chapters 24



Introduction 26


Results 32


Eucalypt Weevils 37


Giant Wood Moths 45

Case Moths 48

Cup Moths 50

Leaf Bag Worms 53




Mirid Bugs 63

v



Psyllids 70

Leafhoppers 73

Planthoppers 75

Clown Bugs 77

Assassin Bugs 79

Ladybird Beetles 81

Praying Mantids 83

Lacewings 85

Discussion 87



Introduction 91


Results 101










Discussion 130



SEASON 134

Introduction 134


vi

Results 153

Discussion 184



Introduction 191


Results 198

Discussion 216



Introduction 221


Results 231

Discussion 237


Important Pests 242







Future Research 257


8 REFERENCES 259

vii

Abstract
























viii


























ix











1























2


Inventory 2007)























3














4

























5
























6











Grafton

Bundaberg

Brisbane

Rockhampton

Coffs Harbour

Urbenville




QLD-NSW

Border


7






















8



Defining Stress





















9

























10

























11

























12





al 2006)
















groups (Table 11)

13


Borers






Sauvard 2004


Abies spp

Larix spp

Pinus spp

Picea spp













Christiansen 1992



14

Borers continued



Eucalyptus spp

Acacia spp





Duffy 1963

Hanks et al 1999




Pinus banksiana

Pinus strobes

Picea abies





Lavallee 1994


Larix decidua

Picea spp

Pinus spp




Schwerdtfeger 1929

Avtzis et al 2000

Kirisits 2004


Pinus sp

Picea sp

Europe

North West Africa

Northern Asia







15

Defoliating Insects






Miles et al 1982


Quercus spp

Tsuga canadensis

Europe

Asia

North America





Davidson et al 1999


Cedrus spp

Larix spp

Pinus spp




Rouault et al 2006

Buffo et al 2007


Quercus spp

Acer spp

Betula spp

Fagus spp

Populus spp





Gasow 1925

Schwerdtfeger 1971

Larsson et al 2000


16

Other Insects





White 1969

Miles et al 1982

17

























18
























19










Wargo 1996

Metaliaj 2003


Fagus silvatica

and Quercus spp



Hendry et al 1998









Barr 1972


Zhonghua et al 2001


20



Phoenix dactylifera

Saudi Arabia

Iraq




Suleman et al 2001



Quercus spp

Castanopsis spp

Acer spp

Rhus spp

Typhina spp

Carya ovata

Europe

Asia

Africa

North America





Shear et al 1917

Hepting 1974

Anagnostakis1984




Ross 1964




Britain



Gibbs et al 1997


Acer spp

Populus spp




Christensen 1940

Bertrand 1967

Jones 1985

Guyon et al1996

21



Australia




Shearer et al 1987

Old et al 1990


Quercus spp

Juniperus spp

Fraxinus spp

Eucalyptus spp

Europe

South America

North America




Luque et al 2002


Pinus spp

Picea spp

Abies spp




Birch 1937

Laughton1937

Eldridge 1961

Lűckhoff 1964

Buchanan 1967



Barker 1979

Gibson 1980

Brown et al 1981

Chou 1982

Swart et al 1985

22



North America

Australia

South Africa




Schoenweiss 1975

Davison 1982


Walker et al 1985

Roane et al 1986

Old et al 1990

Gryzenhout 2006



Quercus spp

Castanea spp

Populus spp






Valentini 1994

Neofusicoccum ribis


E dunnii

E grandis

E camaldulensis

E radiata

E cladocalyx

E marginata

Corymbia calophylla




Shearer et al 1987

Old et al 1990

Luque et al 2002

Slippers et al 2004

23



Southeast Asia

Africa



Surico et al1996

Moricca 2002


Aspen spp

North America





Bier 1939

Thomson 1941




Abebe and Hart 1990

Maxwell 1997

Xylella fastidiosa


and Citrus spp



Boyer 1995

24





development


southern Queensland


Queensland region



plantation age)





pathogens



Thesis Chapters




25






Southern Queensland


Southern Queensland






Plantations


26


Introduction











Stone 2001)











27


























28


























29




1985 Farr 2002)











development

Chapter Aim





presented

30


Site Selection




South Wales






grandis)

Sampling Regime











31

















32

Results










P

P


Oxyops sp

Defoliator

Defoliator

Medium Low

P

P

P

P

P

P

P

P


Paropsis obsoleta

Paropsis variolosa



Paropsisterna sp

Cryptocephalus sp

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

Defoliator

High

Low

Low

High

Low

Low

Low

Low



P

P


Defoliator

Defoliator

Low

Low










P

P


Cardiaspina sp

Sap-sucker

Sap-sucker

Medium

Low



P

P


Amorbus sp

Sap-sucker

Sap-sucker

Low

Medium








33














34

Longicorn Beetles

Order Coleoptera

Family Cerambycidae















2-1B amp C)







(Lawson et al 2002)

35












eventually die

Threat to Industry






Lawson et al 2002)






36





37

Eucalypt Weevils

Order Coleoptera
























38










Threat to Industry













39


40

Chrysomelid Beetles

Order Coleoptera
























41




Biology and Ecology











Symptoms and Damage




Threat to Industry






42















43


44


45

Giant Wood Moths

Order Lepidoptera

Family Cossidae























46

















Threat to Industry






47


48

Case Moths

Order Lepidoptera

Family Xyloryctidae





















49

Threat to Industry








50

Cup Moths

Order Lepidoptera

Family Limacodidae









Common 1990)














51





Symptoms and Damage



Threat









minor pests

52


53

Leaf Bag Worms

Order Lepidoptera

Family Psychidae






















54















Threat to Industry










55




56


Order Lepidoptera

Family Tortricidae





















57






Threat to Industry










Queensland

58


59

Leaf Blister Sawfly

Order Hymenoptera



















1984)





60


Threat to Industry








61

Eucalypt Gall Wasps

Order Hymenoptera























62


Threat to Industry









Queensland


63

Mirid Bugs

Order Hemiptera

Family Miridae

Species Rayieria sp












mimics

Symptoms and Damage




Threat to Industry




64








65

Brown Scale Insects

Order Hemiptera

Family Eriococcidae















1971)







66






Threat to Industry




Wales









67


68


Order Hemiptera

Family Pentatomidae






















69

Threat to Industry




references







closely


70

Psyllids

Order Hemiptera

Family Psyllidae























71


1974)

Threat to Industry















72


73

Leafhoppers

Order Hemiptera

Family Eurymelidae








visible dorsally












of insects occur

74

Threat to Industry










75

Planthoppers

Order Hemiptera

Family Flatidae

Species Siphanta sp















Threat to Industry





76


77

Clown Bugs

Order Hemiptera

Family Coreidae


















2-18B)




78

Threat to Industry





1998)






79

Assassin Bugs

Order Hemiptera

Family Reduviidae
















Role in Plantations






80


81

Ladybird Beetles

Order Coleoptera











(Figure 2-20B)







Role in Plantations




82


83

Praying Mantids

Order Mantodea










Rentz 1966)







Role in Plantations




84


85

Lacewings

Order Neuroptera










D) (Riek 1970)













86

Role in Plantations






87

Discussion























88





















Urquhart 1995)





89

























90






















91


Introduction






















92





Pegg et al 2005)


















(Cheah 1977 Fry 1983)



93














2003)












94


























95



et al 2004)

Chapter Aim






Site Selection







Sampling Regime





Sampling Method


96















Fungal Isolation










97













dark at 20C












98














1990)











99
















Phylogenetics









100






















101

Results










Figure 3-12)


102


Saprophytic fungi




Aspergillus sp

Cladosporium sp

Epicoccum sp

Fusarium sp

Mucor sp

Penicillium sp

Pestalotiopsis sp


Phoma glomerata

Phomopsis diaporthe

Nigrasporum sp

Trichoderma sp












103






Primary Pathogens
















104


Primary Pathogens










105



Field Symptoms




(Figure 3-2A amp B)

Growth on Agar









Ecology and threat







106






plantation industry

107


108


Host E dunnii

Field Symptoms





Growth on Agar











spore (Figure 3-3G)

Ecology and Threat




109








110


A

B

111


Host E globulus

Field Symptoms





Growth on Agar










Ecology and Threat






112




113


Host E grandis

Field Symptoms





Growth on Agar









Ecology and Threat






114



115


Host E globulus

Field Symptoms





Growth on Agar










Ecology and Threat






116




117


Host E dunnii

Field Symptoms




ostioles

Growth on Agar











Ecology and Threat



Queensland

118


119

New Fungal Species















(Table 32)

120







MUCC467 EU300999

MUCC468 EU301000

MUCC649 DQ240133

MUCC693 EU301002

MUCC694 DQ240169


























MUCC668 EU301011

MUCC669 EU301014

































5 changes

100

67

100

100

76

100

75

96

100

55

100

100

100

99

92

85

87

57

100100

99

100

100

100

100

97

100

100

86

98

66

97

91

97

100

84

84

99

98

88

52





121



















122

























123








124


125

















3-11 E)




ex-type MUCC649)




126






127













(Figure 3-12 K)









MUCC648)



128




129


130

Discussion







species

















Asci

131


















132













conditions






glasshouse






133















Conclusion







134


Introduction





















135

























136

























137

















Chapter Aim








138




Site Selection





















139


Size (ha)














E

S

N

W




140

Group - Age

Size (ha)








Fertiliser History

















141

















defined (Table 43)

142



Foliar Yellowing



Foliar Reddening



143





Total Fungal Damage



144









145





Mirid Damage



146


Psyllid Damage






147





Weevil Defoliation




148


Foliar Wasp Galls



Scale Insect Damage



149





150











of each plantation






151








the average leaf






152







plantation groups
















153


















Results

Damage Averages






154



recorded










Mirid Damage 29 9th







Total 100










155

Plantation Age


34 60 43 53

Local Climate


37 58

Seasons


41 30 59 60















South-4 01 0660 01 0231 01 0179








156













North-3



South-1

South-2

South-3

South-4

North-1

North-2

North-4

North-3

157










158



1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th

13th











1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th

13th

14th

15th











159







Table 412)

160



mean 23 18 1 34 14

plusmn SE 14 1 07 44 19

mean 3 3 13 19 16 16 01

plusmn SE 13 13 08 37 13 13 02

mean 19 16 08 28 04 04 01

plusmnSE 16 11 07 45 09 09 02

mean 19 16 08 28 04 04 01

plusmnSE 08 08 1 46 02

mean 22 2 12 27 05 05 04

plusmnSE 13 12 09 42 1 1 11 01

mean 19 16 08 28 04 04


mean 184 183 116 28 28

plusmn SE 6 59 58 41 41

mean 121 12 121 125 04 04

plusmn SE 57 57 67 354 06 06

mean 145 13 25 13 03

plusmn SE 29 21 46 35 05

mean 116 111 116 37 03 08 08 01


Plantation mean 152 138 152 43 06 06 13 09 24

Group plusmn SE 79 79 66 37 12 12 23 2 4

mean 519 222 519 33 02 09 3 03

plusmn SE 74 74 68 7 05 27 69 05

mean 191 143 191 53 01 01 16 04

plusmnSE 116 68 116 45 02 02 26 07

mean 421 155 421 56 02

plusmnSE 75 75 53 72 04

mean 321 165 321 46 02 02 04 09 07 06 02

plusmnSE 173 78 173 56 06 06 12 21 35 22 04

mean 152 152 175 313 22 05 05

plusmnSE 45 45 32 17 87 06 06

mean 371 371 124 7 02 02 88 13

plusmn SE 72 72 25 35 04 04 12 35

mean 174 174 9 71 05 05

plusmn SE 58 58 17 14 07 06

mean 245 245 113 41 77

plusmn SE 85 85 41 82 83

mean 235 235 126 88 15 03 03 22 03

plusmnSE 18 18 43 16 96 05 05 69 18


Nov-04

1

2

1

2

3

4

4

Total

1


Total

3

4

Total

1

2

3

2

3

4

Total

AgeEstate month

May-05

Feb-05

Aug-04


161



mean 33 13 26 38 378 01

plusmn SE 28 28 25 16 165 04

mean 16 154 141 163 32 179 177 145 19 01

plusmn SE 52 48 46 85 17 5 5 46 37 04

mean 135 13 155 222 141 93 92 146 16 01 05

plusmn SE 68 68 52 179 15 4 4 37 35 02 08

mean 343 341 145 43 94 67 65 38 39

plusmn SE 64 64 45 137 18 48 48 5 4

mean 168 159 117 26 229 85 83 83 18 01

plusmnSE 13 13 67 183 185 76 75 75 34 02 05

mean 9 9 55

plusmn SE 55 55 09

mean 9 9 15 44 24 23 147

plusmn SE 38 38 72 73 24 24 69

mean 29 29 75 25 3 3 02 13

plusmn SE 51 51 1 46 22 22 07 11

mean 172 171 92 29 67 67 01

plusmn SE 22 22 44 76 39 39 02

Northern mean 139 139 93 24 29 29 38 03


Group mean 238 238 5 06 58 58 11 163 06

plusmn SE 25 2 18 72 72 18 132 18

mean 35 35 15 63 78 77 26 01

plusmn SE 113 113 34 92 57 55 4 02

mean 34 34 111 17 17 16 14 02 01

plusmn SE 125 125 43 29 09 09 24 04 02

mean 356 356 8 13 05 05 03

plusmn SE 57 57 2 23 07 07 07

mean 321 321 86 25 39 39 03 47 02 04 01

plusmnSE 99 98 37 53 53 53 1 95 09 13 04 02

mean 17 17 193 16 09 03 145 47 38 12 03

plusmn SE 43 43 56 31 27 07 8 14 58 2 09

mean 443 443 12 3 04 04 15 04

plusmn SE 33 33 2 3 09 05 11 04

mean 36 36 16 2 16 01 13 03

plusmn SE 62 62 17 28 35 03 19 09

mean 464 464 13 16 03 02

plusmn SE 87 87 27 23 04 04

mean 359 359 133 21 07 03 36 41 09 04 02

plusmnSE 131 131 49 27 22 05 02 74 83 32 11 05


Total

Total

2

Nov-04

1

2

1

2

3

4

3

4

1

2

3

4

Total

1

3

4

Total


month AgeEstate

May-05

Aug-04

Feb-05


162




















163



















164

Plantations Estates


1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th





Climate Averages







(Figure 4-6C amp D)









165








0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

Mean Rainfall (mm)

Mean Max Temp (C)

Mean Min Temp (C)


Months Months


0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300

Mean

Rain

fall (

mm

)

0

5

10

15

20

25

30

35

May Ju

nJu

l

AUG

SEP

OCT

NO

VDEC

JAN

FEBM

AR

APR

MAY

Months

Tem

pera

ture

(C

)

0

50

100

150

200

250

300M

ean

Rain

fall (

mm

)

Mean Rainfall (mm)

Mean Max Temp (C)

Mean Min Temp (C)

START OF SURVEY

C D

2004 2005 2005 2004

A B

166










P R P R P R P R

Aug 04

Nov 04 01 0438

Feb 05 01 0631 01 0547

May 05 01 091 01 0934 01 077













167







Four Groupings

Stress 017

168

Rank

Seasons


1st

2nd

3

rd

4th

5th

6th

7th

8th

9th

10th

11th

12th










169















plantation group




170


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










171


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










172



















categories






173


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










174


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










175
















176



00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D


G H










177


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H











178





Damage category







179


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










180




00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










181

Mirid Damage












182


00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

-100

00

100

200

300

400

500

600



erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600



Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

E F G H










183
























less than 1)

184

Discussion

























185








and season


















186













infestations













187









group











trees






188





















Humphreys 1965)





189














dieback












190






Economic Impacts









191


Introduction












and Wingfield 1996)











192



Queensland)
























193








(Dungey et al 2000)

Chapter Aim
















194









195


E dunnii





E grandis

(Flooded gum)




E globulus

(Blue gum)




E tereticornis

(Forest red gum)






196


E camaldulensis

(River red gum)




E urophylla





Hybrid taxa













197



Sampling Regime


Climate Data



and Figure 4-6)








198











those differences

Results









199













Total 100

Comparing Taxa















damage



th) caused by


200





201






Pure Taxa


1st

2nd

3

rd

4th

5th





Hybrid Taxa


1st

2nd

3

rd

4th

5th

6th








202















G1 G2

G3

G4

203










2004



P R P R P R P R

Aug 04

Nov 04 01 0763

Feb 05 01 0634 01 0271

May 05 01 0757 01 0481 01 0562





(Table 56)

Rank

Seasons of Sampling


1st

2nd

3

rd

4th

5th







204
















205












206

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

un

E g

lob

E g

ran

E teret

E g

ran x cam

E teret x uro

E u

ro x cam

E u

ro x gran

Eucalypt Taxa

Pe

rc

en

tag

e o

f D

am

ag

e (

)



A B C D

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)



A B C D

207






Total Fungal Damage








208

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)



A B C D

00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u


Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)



A B C D

209

Foliar Yellowing












210



00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt TaxaP

erc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)A B C D



00

100

200

300

400

500

600

700

E du

nE

glob

E gra

nE

teret

E gra

n x ca

m

E te

ret x uro

E uro

x camE

uro x gra

n

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

00

100

200

300

400

500

600

700

E d

unE g

lobE g

ranE teret

E g

ran x cam

E teret x uro

E u

ro x camE u

ro x gran

Eucalypt Taxa

Perc

en

tag

e o

f D

am

ag

e (

)

A B C D

211





212

Aug-04

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 92 196 97 326 264 313 292 149

plusmn SE 07 14 17 34 00 23 47 21


M 92 196 94 326 250 313 292 149

plusmn SE 60 60 42 42 42 42 42 42


M 132 120 88 163 183 219 198 167

plusmn SE 17 13 19 24 23 18 21 23


M 09 172 14 00 00 00 00 00

plusmn SE 21 201 24 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Fungal Damage

M 144 150 94 288 83 167 410 128

plusmn SE 17 11 28 45 29 15 37 19

Foliar Yellowing

M 174 00 139 56 42 00 56 111

plusmn SE 63 00 77 56 42 00 56 77

Scale Insect Damage

M 00 00 03 00 00 00 00 00

plusmn SE 00 00 05 00 00 00 00 00


213

Nov-04

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 264 135 205 178 104 188 354 128

plusmn SE 19 12 21 29 07 09 51 08


M 264 135 25 177 14 188 354 128

plusmn SE 60 60 42 42 42 42 42 42


M 214 384 125 104 83 125 208 87

plusmn SE 15 25 00 14 14 00 51 07


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 51 00 00 00 00 00 00

plusmn SE 00 100 00 00 00 00 00 00

Total Fungal Damage

M 00 121 42 21 00 00 111 00

plusmn SE 00 34 23 07 00 00 51 00

Foliar Yellowing

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


214

Feb-05

E dunnii

E globulus

E grandis

E tereticornis






M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Total Defoliation

M 373 167 146 417 125 458 436 413

plusmn SE 22 12 07 34 05 24 25 30


M 373 161 146 417 125 458 431 413

plusmn SE 60 60 42 42 42 42 42 42


M 175 239 146 153 125 167 156 125

plusmn SE 08 20 07 13 00 15 17 00


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


M 00 00 03 00 00 00 00 00

plusmn SE 00 00 06 00 00 00 00 00

Total Fungal Damage

M 30 00 10 340 00 292 188 264

plusmn SE 08 00 06 43 00 15 37 47

Foliar Yellowing

M 36 00 28 00 00 00 14 56

plusmn SE 29 00 19 00 00 00 14 33

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


215

May-05

E dunnii

E globulus

E grandis

E tereticornis






M 307 416 340 00 07 63 10 00

plusmn SE 66 65 85 00 05 16 11 00

Total Defoliation

M 429 117 219 244 66 417 549 444

plusmn SE 22 12 37 21 04 15 27 15


M 429 116 219 243 66 417 549 444

plusmn SE 60 60 42 42 42 42 42 42


M 98 70 101 69 63 63 267 163

plusmn SE 08 03 15 05 00 00 38 27


M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00

Foliar Wasp Galls

M 00 00 00 00 00 00 00 111

plusmn SE 00 00 00 00 00 00 00 192


M 02 00 00 00 00 00 00 00

plusmn SE 04 00 00 00 00 00 00 00

Total Fungal Damage

M 73 193 49 03 00 00 10 10

plusmn SE 21 23 27 04 00 00 06 06

Foliar Yellowing

M 00 00 00 14 00 333 83 42

plusmn SE 00 00 00 14 00 123 58 23

Scale Insect Damage

M 00 00 00 00 00 00 00 00

plusmn SE 00 00 00 00 00 00 00 00


216

Discussion
























217












McGrath 1999)














218



Comparing Taxa



















oviposition of eggs



219









Conclusion

















220














221


Introduction






















222


























223























and Mohammed 1999)



224





















Chapter Aim




225






expression





tissue



226
























227




(White et al 1990)






Site Selection















228


















229



















230


1B Control
















experiments

Measuring Lesions











231







Statistics




treatment

Results


either produced a






232















W

L

233

64)











1A Control 20

2A H eucalypti 40

3A N ribis 40








00

1000

2000

3000

4000

5000

6000

Control

H eucalypti

B ribis

C eucalyptic

ola

H eucalypti +

B ribis

H eucalypti +

C e

ucalypticola

B ribis + C

euca

lyptic

ola

H eucalypt +

B ribis

+ C e

ucalypticola

Treatments

Mean

Lesio

n S

everi

ty I

nd

ex (

len

gth

x w

idth

) m

m LSD (5) = 12210


1A

2A

3A

4A

5A

6A 7A

8A

234











1B Control 20

2B 1 H eucalypti 90


4B 3 H eucalypti 80

5B 1 N ribis 95

6B 2 N ribis 80

7B 3 N ribis 50

8B 4 N ribis 70

9B 5 N ribis 70






235























0

50

100

150

200

250

300

350

400

450

500

Con

trol

1 H

euca

lypt

i

2 H

euca

lypt

i

3 H

euca

lypt

i

1 B

rib

is

2 B

rib

is

3 B

rib

is

4 B

rib

is

5 B

rib

is

1 C

euca

lypt

icol

a

2 C

euca

lypt

icol

a

3 C

euca

lypt

icol

a

4 C

euca

lypt

icol

a

Isolate species

Lesio

n S

everi

ty I

nd

ex (

len

gth

x w

idth

)

mm

LSD (5) = 7500

1B

2B 3B

4B

5B

6B

7B

8B 9B

10B 11B

12B 13B

236














treatments

Fungal Species


00

1000

2000

3000

4000

5000

6000

Cont

rol

H e

ucalyp

ti

B r

ibis

C e

ucalyp

ticola

Fungal species

Mean

Lesio

n S

everi

ty I

nd

ex (

len

gth

x

wid

th)

mm


Neofusicoccum ribis


Winter

Summer

237








treatment




Discussion













238


























239

























240


rainfall










Conclusion













241



242


Important Pests






















243

Important Pathogens
















al 2005)







high rainfall)

244

Economic Impacts

























245







future
















drought


246










al 2002)















247

inciting factors











248








illustrated





249








250




251















Threshhold

Pivot Threshhold

Pivot

Threshhold Pivot

252


















Threshhold Pivot

Threshhold Pivot

Threshhold Pivot

253




as follows




















254



deficiency



1 Drought Impacts


















255
























256







4 Sampling Regime










5 Weather Data







257

more efficient

Future Research















and adults






258

2 Leaf Pathogens






3 Canker Pathogens








Concluding Remarks






259

8 References






74






85-110








1-44








260









13 197-205






















261






167














346






155-163





106

262













Research 49 153-174













49 239-42





263






1318-1320





101 1060-1064





















264






173





370














Canberra






Japan

265











Ottawa



Entomology 9 65-72













Washington DC





266



44




Management 54 69-87























267







21 (2) 53-63
























268


London















Sciences 12 438-441















269



10 537-586


Plymouth UK






















15 409-416


no 74109rsquo

270






97



278



Paul Minnesota USA












441-458





532



271








HMSO London


10-11



















272



Science 63 597-612


















Bulletin 4 41-94











273



Workshop





Science 199 941-945







95 551-557


Mycology 41 339-339





Wales Beecroft NSW









274




of Ecology 12(2) 125-138










Dilemma FAO Rome








Zoology 28 83-90




23(4) 497-507



Science 21 185-193

275







25





Research 94 313-319








33 (2) 111ndash114









276




Publishing











Forestry 28 394-411












Forestry 91 215-221





277


Management 26 38-40





series No 4













Mycology 55 35-52







35-52





278




10-11





The Netherlands













153-162









279



353ndash361






565


1 31-50


67





Extension Nevada


Press Inc Hong Kong


Melbourne









280





















53







ACT



281












119










258 CSIRO Australia










282




















Netherlands





York





283



129-147




















277-83







284















No 982727 pp 28-38


London Academic














285




165-185



















19(1) 7ndash13









286









399 p






















287







Forestry 37 32-42














Management Perth











288




11-18














Science 33 362-372


28(1) 8-10


Australia 28(2) 19









289
















Forestry 69 270-274















290













New Zealand No 57



47-57















291




















Parana Brazil












292







101-115








Botany 29 675-88















293












Botany 46 735-742











138 23-31








294
























43(1) 95-124







295



427ndash33



























27-39

296






















113-122



Disease 71 140-145




Mycologist 19 57-62

297







New York




Mycology 50 343-358



1031ndash1034




Water Queensland



365-369




New York






298



















81









Australia




299









and Water Canberra


















Disease 76(7) 744-746




300

























33 109-110




Botany 54 543-545



301








Beecroft NSW








32 295-402









8(2) 155-160



Genetics 43 366-372




302













23 350-370











CSIRO Melbourne



29 199-208



303



Press Boca Roton





Press







202 67-82









440







304











1493






University





Botany 62 325-327








239-257

305











Press)



Queensland






Tokyo Tokyo











Urumqui P R China

306








84




















Hydrology



Disease 85 745-749

Documents

The Ecology of Insect Pests and Fungal Pathogens of Drought