Download role of insects in forest ecosystems

Lecture Notes: Forest Insects
ENT/PLPATH/FWE500
Insects and Disease in Forest Resource Management
Kenneth F. Raffa
Dept. Entomology
345 Russell Laboratories
University of Wisconsin
Madison, WI 53706
262-1125
[email protected]
http:entomology.wisc.edu/~ raffa
TABLE OF CONTENTS
Role of Insects in Forest Ecosystems
.............................................. 4
Introduction to Entomology
.............................................. 6
Classification and Diversity of Insects
.............................................. 6
Insect Structure & Function
.............................................. 6
Growth & Development
.............................................. 7
Major Insect Orders
.............................................. 7
Insect Behavior
.............................................. 10
Population Dynamics
..............................................
Role in Forest Entomology
..............................................
Population Processes
..............................................
Population Models
..............................................
Life Tables
..............................................
Population Distribution
..............................................
Insect Population Sampling Techniques ..........................................
11
11
11
11
14
15
15
Principles of Integrated Pest Management. ............................................. 16
Economic Injury Levels
.............................................. 16
Available tactics of Forest Pest Control..............................................
Integrated Pest Management
.............................................. 17
Root Insects
.............................................. 18
Introduction
.............................................. 18
Impact on Stand
.............................................. 18
Management
.............................................. 19
Conifer Root Weevils
.............................................. 19
Bark beetles, White grubs, Invasive weevils, Lepidoptera, Homoptera
Stem Colonizing Bark Beetles
Introduction
Biology
Major Species
Management of Bark Beetles
16
21
.............................................. 22
.............................................. 22
.............................................. 22
.............................................. 22
.............................................. 24
Wood Borers
.............................................. 26
Introduction
.............................................. 26
Native Species
.............................................. 26
Metallic Borers, Long-horned Beetles ..............……............ 26
Horntails, Lepidoptera
.............................................. 27
Ambrosia Beetles
.............................................. 28
Invasive Species
.............................................. 29
Emerald Ash Borer
.............................................. 29
Asian Longhorned Beetle
.............................................. 29
Pine Sawyer / pine wilt nematode ......................................... 29
2
Horntail: Sirex noctilio
.............................................. 30
3
Tree Response to Defoliation
.............................................. 31
Introduction
.............................................. 31
Physiology of Defoliation
.............................................. 31
Relation of Control Options to Life History of Defoliating Insects ....32
Hardwood Defoliators
Lepidoptera
Coleoptera
Diptera
Orthoptera and Phasmodea
Thysanoptera
Diptera, Hymenoptera
.............................................. 33
.............................................. 33
.............................................. 36
.............................................. 36
.............................................. 36
.............................................. 37
.............................................. 37
Conifer Defoliators
Budworms
Tussock Moths
Caseworms and Bagworms
Conifer Sawflies
Regionally important defoliators
..............................................
..............................................
..............................................
.............................................
..............................................
…..........................................
38
38
39
40
40
41
Fluid Feeding Insects
Introduction
Homoptera
Mites: Acarina
Other
..............................................
..............................................
..............................................
..............................................
..............................................
42
42
42
44
44
Shoot and Tip Insects
Introduction
Weevils
Shoot Moths
Bark Beetles
..............................................
..............................................
..............................................
..............................................
..............................................
45
45
45
46
47
Cone and Seed Insects
Cone Moths
Seed Moths
Cone Beetles
.............................................. 48
.............................................. 48
.............................................. 48
.............................................. 48
Gall Insects
.............................................. 49
Appendix 1: Insect Population Sampling Techniques
Appendix 2: Major Categories of Insecticides; Mechanisms of Insecticide Resistance
4
ROLE OF INSECTS IN FOREST ECOSYSTEMS
I.
Herbivory (=Phytophagy)
A.
Host Range: Mono-, oligo-, poly- phagous
Implications to pest management
B.
Plant parts (highly specialized)
Microsite
C.
Impact
1. Can range from marginally beneficial to lethal
2. Implications to Management:
Stand level losses in forests
Individual losses in ornamentals
3. Beneficial: Bio Control of weeds (Alder) & Fungi
4. Ecological: Affects competition between plants
Can influence succession, forest structure, gaps, fire
5. Vector plant pathogens
Insect mouthparts
Sometimes mechanical, sometimes quite specialized
6. Must consider at multiple levels
Plant part: Tree may compensate: Diversion of energy, abscission
Stand level: Increase of light, moisture, release from competition
Release of bound biomass
II.
Predation & Parasitism
A.
Host range
B.
Specialize on insect stages: Eggs, Larvae, Pupae, Adults
C.
Sophisticated prey searching mechanisms
Odor - Plant wounds, insect
D.
Management: BioControl
5
Ecology: Can enhance diversity of ecosystem; Not allow any one herbivore to
reach high densities
III. Food For other Animals
A.
Food webs
B.
Antipredator mechanisms
Camouflage
Mimicry: Toxic insects, wasps, etc.
Chemical Defenses
Implications to Management: Sampling
Biological Control
IV. Saprophagy
Importance
Context of pest: Termites, Carpet beetles
V.
Pollination
2/3 of flowering plants
Forestry - mostly hardwoods
VI.
Pest Actions of Insects.
Only “pest” because they interfere with human objectives
Major pests include both introduced and native
6
INTRODUCTION TO ENTOMOLOGY
I.
CLASSIFICATION AND DIVERSITY OF INSECTS
A.
Arthropods
Phylum Arthropoda: Key Characteristics
Exoskeleton
Segmented body parts
Paired appendages
Open circulatory system
Ventral nerve cord (decentralized)
Open respiratory system
Crustacea - Aquatic - 2 prs antennae - 2 body parts (cephalothorax, abdomen)
Millipedes - more than 1 pr/segm, scavengers
Centipedes - 1 pr/segm, predators
Arachnids - Spiders (pred), Ticks, Mites (Some are important pests) no antennae, 4
prs legs,
B.
Insects
Class Insecta: Key Characteristics 3 distinct body parts
3 prs legs
2 pr wings
1 pr antennae
Most diverse of all groups - 1 - 3 million species - @90% of all animals, over half of
all life forms
C.
Useful Field Guides to Forest Insects and Their Damage
Drooz: Eastern Forest Insects, USDA Misc. Publ. 1426, 1985
Furniss & Carolin: Western Forest Insects, USDA Misc. Publ. 1139, 1977
II.
INSECT STRUCTURE AND FUNCTION
A.
Exoskeleton
B.
Major Body Regions
C.
Nervous System
D.
Circulatory System
E.
Respiratory System
7
III.
F.
Digestive System
G.
Reproductive System
GROWTH AND DEVELOPMENT
A.
Types of Development
1. Hemimetabolous ‘Incomplete metamorphosis’
2. Holometabolous ‘Complete metamorphosis’
B.
Moulting
C.
Endocrine control
1. Ecdysone
2. Juvenile Hormone
D.
Life History Stages
1. Diapause
2. Dormancy:, Aestivation
3. Diurnal Rhytms
4. Migration
IV. MAJOR INSECT ORDERS
Introduction
Phylogenetic Order
28 orders; 8 orders contain >95% of species
A.
Orthoptera (Grasshoppers);
Related orders include Phasmatodea (walking sticks) and Mantodea (mantids)
1.
2.
3.
Chewing mouthparts
Incomplete metamorphosis
Leathery forewings, Membranous hindwings
Most not very important in forestry
Some important groups:
Walking sticks on oak in US; Eucalyptus in New Zealand
Grasshoppers on oak
B.
Blattodea (formerly Isoptera) - termites
1. Chewing mouthparts
2. Incomplete metamorphosis
8
3. Highly social
Very common in forests; feed on cellulose (Indigestible - require protozoa and
bacteria - basis of their social systems)
Beneficial - nutrient cycling
Damaging - wood products, sometimes trees
Blattodea - roaches
C.
Thysanoptera - Thrips
1. Extremely small - Feathery wings
2. Highly modified mouthparts:
Intermediate between chewing & sucking
Piercing & sucking
Asymmetric
3. Intermediate metamorphosis
Several important species in forestry but not many
D.
Hemiptera - Bugs
1. Sucking mouthparts
2. Incomplete metamorphosis
3. Suborders
a. Heteroptera
Some plant pests, many predators, some human pests
b. Homoptera - Aphids, Scale insects, Spittle bugs, cicadas
Plant feeders
Sometimes quite complex development
Important ectors of plant pathogens
E.
Coleoptera
1. Largest insect order (@300,000): Most important pest group in forestry
2. Chewing mouthparts
3. Complete metamorphosis
4. Elytra (Thick & horny; Cover hindwings in flight)
Highly Diverse: Plant feeders, predators
Immatures - Grublike, Some free living
F.
Diptera - flies – Fourth large orderest
9
1. Sucking mouthparts - adults
2. Complete metamorphosis
3. Adults - 2nd Pair of wings highly reduced "Haltere" – Stabilizing organs
4. Larvae - maggot like
Not major pest of forests
Important vectors of pathogens causing Human Disease
Parasitic flies - Tachinidae
Look like houseflies - hairy
Female attaches egg to larva (mostly Lep) or leaf
Predacious flies - Asilidae (Robber flies); Dolichopodidae
G.
Lepidoptera
1. Third largest insect order: Second important pest group in forestry
2. Sucking mouthparts
3. Complete metamorphosis
4. Adults - Scaly wings
5. Less diversity than Beetles: Almost all are plant feeders as larvae.
Most feed on nectar as adults (some do not feed)
6. Larvae are damaging stage "caterpillars" - 3 pairs of true legs
4 - 5 pairs fleshy prolegs
7. Types of feeding
Defoliation, Mine leaves, stems, buds
H.
Hymenoptera - Wasps, Bees, Ants, Sawflies, Woodwasps, Parasitic wasps
1. Second largest insect order; Most beneficial group in forestry
2. Chewing mouthparts - Modified
3. Complete metamorphosis
4. Adults - Membranous wings;
5. Ovipositor - Highly specialized - Defense, Insertion into plants or insects
6. Three Important Ecological Groups
a. Plant - feeders: Sawflies, woodwasps, galls: Pests
b. Parasites - Ichneumonidae & Braconidae: Benefit
c. Social Insects - bees, ants, wasps - pollination, predation:
7. Larvae
a. Caterpillar like (Sawflies > 5 prs prolegs)
b. Grublike
V.
INSECT BEHAVIOR
A.
Introduction: "Observable Physiology"
10
B.
C.
D.
Feeding Behavior
1.
Types of feeding
2.
Host range
3.
Specialization on plant parts
4.
Host location
Reproductive Behavior
1.
Mate location
2.
Mating frequency
3.
Oviposition
Orientation
11
POPULATION DYNAMICS
I.
II.
ROLE IN FOREST ENTOMOLOGY
A.
Prediction
B.
Management
B.
Characteristics of Populations
POPULATION PROCESSES
A. Birth, Death, Immigration, Emigration
III.
POPULATION MODELS
A.
Exponential Growth
dn/dt = rN
Nt = N0ert
B. Sigmoid Growth
Carrying Capacity K
dn/dt = rN {(K-N)/K}
Nt= K/ (1+B-rt)
C.
Competition
D.
Predation
E.
Density Independent and Density Dependent Factors
Density Independent Factors: The proportionate effect on population processes
(Birth, death, reproduction, movement) is not related to population density.
12
1. Example: Early frosts in 1995, 1999, & 2007 caused high mortality to a budworm. But
the proportionate mortality is not related to population density
Population
After frost
1000
500
800
No frost
50000
No frost
75000
37500
100
No frost
200
100
Yr.
2002
2003
2004
2005
2006
2007
Before frost
Mortality
500
% Mortality
50
37500
50
100
50
2. Example: Mortality and % mortality vary; Proportionate mortality related to
precipitation, but not density.
Year
2003
2004
2005
2006
2007
Population
May 1
June 1
500
100
2000
1000
1500
600
1000
600
2500
800
Precipitation
7
4
5
3
6
Mortality
400
1000
900
400
1750
% Mortality
80
50
60
40
70
100
50
25
3000
2500
2000
1500
1000
500
0
0
% Mortality
75
2
3
4
5
6
7
Precipitation
Population
13
8
F. Density Dependent Factors: The proportionate effect on population processes (Birth,
death, reproduction, movement) is related to population density.
1. Example:
Population rates and mortality due to disease are listed below. As the population
increases, both the absolute mortality and the % mortality increase
Yr.
2002
2003
2004
2005
2006
2007
Population
1000
5000
100000
800
10000
50000
Mortality
10
500
90000
8
2000
25000
% Mortality
1
10
90
1
20
50
100
% Mortality
75
50
25
125000
100000
75000
50000
25000
0
0
Population
Types of density-dependent forces
1.
Competition
a. Starvation
b. Can sometimes be have Non-lethal Effects:
Examples: Delayed Development, Reduced Fecundity
2.
Predation and Parasitism: Both Numerical and Functional Responses
a. Numerical response: Predator population increases when prey abundant
b. Functional response: Consumption / predator increases when prey abundant
14
3.
Disease
4.
Density Dependence Can Sometimes be Positive
a. Allee Effects: Significance to invasive species
5.
C.
D.
IV.
Some environmental effects can have a density-dependent component
Carrying Capacity, Sigmoid Growth and Equilibrium Behavior
1.
Effect on Population and Per Capita Growth Rates
2.
Time delays
3.
Integrating Density-Dependent and Density Independent factors
Genetic Change
LIFE TABLES
Life tables present the number of surviving individuals from an initial cohort, at a series of
time intervals. They are used to predict future population densities, both between and
within generations.
A.
Age-Specific death rate: percentage that dies during a particular stage.
B. Cumulative death rate: percentage of original cohort that dies before a particular life
stage is reached.
Example:
Stage
Egg
Larva - 1
-2
-3
-4
Pupa
Adult
Age - Specific
Death Rate
50%
60%
50%
40%
58%
20%
--
No. Surviving
1000
500
200
100
60
25
20
Cumulative
Death Rate
-50%
80%
90%
94%
97.5%
98%
C. Key Factors
Source of Stage-specific mortality that best predicts year to year fluctuation
Not necessarily the highest mortality factor
15
How is it computed? a) calculate the age specific mortality rates; b) determine what
factor is most responsible for mortality during each stage; c) determine which stagespecific mortality is most correlated with generation to generation change ; d) What
is the agent that causes this mortality?
V.
POPULATION DISTRIBUTION
A.
Space
Random, Clustered, Even
B.
Time
1.
Seasonal abundance
2.
Outbreak behavior
C. Space * Time Interactions
VI. INSECT SAMPLING
A. Sampling vs. Collecting
B.
Absolute vs. Relative Methods
C.
Appendix 1
16
PRINCIPLES OF INTEGRATED PEST MANAGEMENT
I.
II.
ECONOMIC INJURY
A.
Relationship of Population Density to Yield Loss
B.
Sampling
C.
Decisions to Employ Control Measures
AVAILABLE TACTICS OF FOREST PEST CONTROL
A.
B.
C.
D.
Biological Control
1.
Natural Components
2.
Introduced Species
3.
Enhancement of Native Control Agents
4.
Biotic Insecticides
5.
Advantages, Disadvantages
Silviculture
1.
Species Composition and Planting Schedules
2.
Thinning
3.
Harvesting Schedules
4.
Advantages, Disadvantages
Resistance Breeding
1.
Genetic Variation
2.
Operational Considerations
3.
Biotechnology
4.
Advantages, Disadvantages
Exclusion and Eradication
1.
Quarantine
2.
Relating Exclusion to Other Control Measures and Insect Biology
17
E.
Chemical Control
1.
III.
Insecticides (Appendix 2)
a.
Types
b.
Advantages and Disadvantages
c.
LD values
d.
Systemic
2.
Pheromones
3.
Insect Growth Regulators
4.
Antifeedants
INTEGRATED PEST MANAGEMENT
A.
Optimum Combination
B.
Applying General Principles To Specific Situations
18
ROOT INSECTS
I.
INTRODUCTION
A.
Increased importance
B.
Taxonomy of major groups
C.
General features of root insects
D.
II.
1.
Slow development time
2.
Oligophagous
3.
Orientation to host odors
4.
Clustered distributions
5.
Low fecundity
6.
Close symbiotic relationships with fungi
High Level of Niche Partitioning
1.
Host age and condition
2.
Microhabitat
3.
Behavior
IMPACT ON STAND
A.
B.
Direct
1.
Larval feeding
2.
Adult feeding
Indirect
19
III.
IV.
A.
B.
MANAGEMENT
A.
Sampling
B.
Control
1.
Chemical
2.
Biological
3.
Silvicultural
4.
Tree breeding
CONIFER ROOT WEEVILS
INTRODUCTION
1.
Types of Damage
2.
Weevils: Curculionidae
a.
Identification
b.
Description
3.
Relationships With Other Insects and Fungi
4.
Increased Importance
MAJOR CONIFER ROOT WEEVILS:
1.
Pine Root Collar Weevil: Hylobius radicis
a.
Hosts
b.
Distribution
c.
Life cycle
d.
Damage
e.
Stand factors
f.
Control and Management
20
2.
3.
Pine Root Tip Weevil: Hylobius rhizophagus (H. assimilis)
a.
Hosts
b.
Distribution
c.
Life cycle
d.
Damage
e.
Control and management
Pales Weevil: Hylobius pales
a.
Hosts
b.
Distribution
c.
Life cycle
d.
Damage
e.
Control and management
4.
Pitch-eating Weevil: Pachylobius picivorus
5.
Other Conifer Root Weevils
a.
Warren's Collar Weevil
b.
Strawberry Root Weevil
c.
Eastern Pine Weevil: Pissodes nemorensis
d.
Hylobius abietis: Europe
21
V.
BARK BEETLES (COLEOPTERA: SCOLYTIDAE)
A.
Hylastes
1.
Pests of western Douglas fir plantations, vectors of
Black Root Stain disease
B.
VI.
Pseudohylesinus
WHITE GRUBS (COLEOPTERA: SCARABAEIDAE)
A.
Biology
B.
Plantation Pests, Alternate Hosts
C.
Control
VII. INVASIVE ROOT WEEVILS OF NORTHERN HARDWOOD FORESTS
A. General Biology and significance
B. Phyllobius oblongus
C. Polydrusus sericeus
D. Sciaphilus asperatus
VIII.
LEPIDOPTERA
A.
IX.
Conifer swift moth
HOMOPTERA
A.
Aphids
B.
Cicadas
22
STEM COLONIZING BARK BEETLES
I.
INTRODUCTION
A.
II.
1.
Bark beetles
2.
Ambrosia beetles
B.
Economic Importance
C.
Distribution
D.
Classification By Vigor of Hosts Selected
BIOLOGY
A.
Life History
B.
Host Resistance
C.
Pheromones
D.
III.
Taxonomy: Coleoptera: Curculionidae: Scolytinae (formerly Scolytidae)
1.
Aggregation
2.
Anti-aggregation
Microorganisms
1.
Blue Stain Fungi: Ophiostoma, Ceratocystis
2.
Digestion
E.
Relationship of Life Stages to Control Tactics
F.
Factors Predisposing Trees to Attack
MAJOR SPECIES
A.
Mountain Pine Beetle: Dendroctonus ponderosae
1.
Northwest and Rocky Mountains
2.
Lodgepole Pine, Ponderosa Pine, Sugar Pine, White Pine
3.
Grosmannia clavigerum formerly O. clavigera
23
4.
Responses to climate change: Changes in voltinism, habitats, geographic
range
B.
C.
D.
Western Pine Beetle: Dendroctonus brevicomis
1.
California and southwestern Rocky Mountains
2.
Ponderosa pine
Southern Pine Beetle: Dendroctonus frontalis
1.
Southern states
2.
Loblolly, Virginia, Short Leaf Pine
3.
O. minus; Basidiomycetes; Mites
Spruce beetle: Dendroctonus rufipennis
1.
Alaska, British Col., Rocky Mtn states, North Central states, New
England
E.
F.
G.
2.
White spruce, Sitka Spruce, Lutz spruce
3
Variations in population behavior: Region; Population phase
Pine Engraver: Ips pini
1.
Transcontinental
2.
All pines, some spruces
3.
Major Lake States bark beetle
4.
O. ips
European Spruce Bark Beetle: Ips typographus
1.
Europe
2.
C. polonica
Turpentine Beetles: Attack Base of Weakened Trees; Vector Leptographium fungi
1.
Red Turpentine Beetle: Dendroctonus valens
a.
b.
2.
Transcontinental
Introduced into China in mid 1980’s.
Black Turpentine Beetle: Dendroctonus terebrans
a.
Southern States
24
3. Dendroctonus murrayanae in western Canada, high elevations of US Rocky Mts
4. Dendroctonus micans in Europe
25
H.
Other Conifer Bark Beetles - Moderately Aggressive. Capable of outbreaks under
stress conditions
1.
2.
3.
Southern States
a.
Eastern 5 Spined Pine Engraver: Ips grandicollis
b.
Small Southern Pine Engraver: Ips avulsus
Western States
a.
Douglas Fir Beetle: Dendroctonus pseudotsugae
b.
Fir Engraver: Scolytus ventralis
Eastern and Lake States
a.
I.
Eastern Larch Beetle: Dendroctonus simplex
Dutch Elm Disease
1.
Imported Pest Complex
2.
Smaller European elm bark beetle: Scolytus multistriatus, C. ulmi
Native elm bark beetle: Hylurgopinus rufipes
J.
3.
Biology of beetle, vectoring of fungus, and transmission across root grafts
4.
Control
Other Bark Beetles Colonizing Angiosperms
1.
IV.
Hickory Bark Beetle
MANAGEMENT OF BARK BEETLES
A.
Impacts
1. Forest Products
2. High Values Trees
a. Recreation, Watershed
b. Private
3. Wilderness
a. Old Growth
b. Endangered Species
26
c. Dispersal
B.
C.
Chemical
1.
Insecticides
2.
Pheromones
Biological
1.
2.
D.
a.
Predators - Beetles, Flies, Mites, Woodpeckers
b.
Parasites - Wasps, Flies, Nematodes
c.
Disease Agents
Utilizing Biological Control
a.
Problems
b.
Role in Sanitation
Silviculture
1.
2.
3.
E.
Natural Enemy Complex
Augmentation of Host Resistance
a.
Thinning
b.
Site Selection
c.
Harvesting Schedule
Reduce Food Base
a.
Age mosaic
b.
Removal of large trees
c.
Timing of operations
Population Removal
a.
Removal of Infested Trees
b.
Destruction of Slash
Survey Methods and Population Prediction
27
WOOD BORERS
I.
INTRODUCTION
A.
General Biology
B.
Host Location
C.
Ecological Role
D.
Economic Impacts
Native Species
I.
"METALLIC BORERS," "FLAT-HEADED BORERS"
A.
Coleoptera: Buprestidae
B.
Life Cycle
C.
1.
Adult characteristics
2.
Larval characteristics
3.
Larval mines
Ornamental and Forest Pests
1.
Bronze Birch Borer: Agrilus anxius
a.
D.
E.
Resistant Varieties: Japanese White Birch--"Whitespire"
2.
Two-lined Chestnut Borer Agrilus bilineatus
3.
Control
Plantations
1.
Poplar root girdler: Agrilus horni
2.
Turpentine borer: Buprestis
Lumber Yards
1.
Golden buprestid: Buprestis
28
II.
"LONG-HORNED BEETLES," "ROUND-HEADED BORERS"
A.
Coleoptera: Cerambycidae
B.
Life Cycle
C.
1.
Adult characteristics
2.
Larval characteristics
3.
Larval mines
Host relationships
1.
Living trees
a.
Sugar Maple Borer
b.
Red Oak Borer
c.
Locust Borer
d.
Poplar Borers:
1.
e.
III.
Management
2.
Recently felled trees
3.
Old moist wood
4.
Dry seasoned wood
5.
Twig and stem girdlers
"HORNTAILS"
A.
V.
Saperda spp: association with Hypoxylon cankers
Hymenoptera: Siricidae
1.
Description
2.
Life Cycle
3.
Control
LEPIDOPTERA: Cossidae
A.
Oak carpenterworm
29
VI. AMBROSIA BEETLES
A. Coleoptera: Scolytidae (and related families)
1.
Biology
2.
Economic impact
3.
Trypodendron
B. Management
1.
2.
Cutting Operations
a.
Removal of brood trees and logs
b.
Rapid removal of felled trees
Storage
a.
Distance from sources of beetles
b.
Maintain minimum inventories
c.
Water Misting
d.
Pheromones
VI. Termites
VII. Other
A.
Carpenter Ants: Formicidae
B.
Coleoptera: Bostrichidae, Curculionidae
C.
Diptera: Agromyzidae
30
Invasive Species
A.
B.
Emerald ash Borer
1.
Host range, Biology
2.
Geographic range
3.
Means of spread
4.
Control
Asian Long-Horned Beetle
1.
Introduced into NY in 1996; Chicago 1998.
2.
Native range: China, Japan, Korea
3.
Hosts
4.
4.
C.
a
Primary: Maple, poplar, willow
b.
Other: Birch, elm, horsechestnut
Life history
a.
@ 30 eggs/female; hatch in 10 days
b.
Camrial Region: L1-L3; Sapwood: L4-L5.
c.
Pupate in tree
Control: Sanitation, chemical
Pine Sawyer--Vector of Pine Wood Nematode
1.
Japan, Portugal
a.
b.
2.
Causal agents
1.
Native beetle: Monochamus
2.
North American nematode: Bursaphelencus
Host--Japanese red pine
Biology
31
3.
Impact on host: Death of ray and axial paranchyma cells surrounding duct
epithelium, wilt
4.
Control
a.
Insecticides
b.
Sanitation
c. Introduced entomophagous nematode
D.
Sirex noctilio in Australia, New Zealand, S. Africa
1.
2.
Introduced Insect/Fungal Complex, Introduced Tree
a.
Amylastereum spp & Stereum spp
b.
Pinus radiata (Monterey Pine)
Life Cycle
a.
Adults attracted to stressed trees (Monoterpenes)
b.
Insert egg in first drill
Insert fungal arthrospores & mucus in second drill
c.
Host responds by accumulating resins & polyphenols
d.
Fungi & mucus impede host response
e.
Multiple attacks
3.
Wilt Symptoms & Tree Death
4.
Control
a.
Biological
1.
Nematode: Deladenus spp
b.
Sanitation
c.
Resistance Breeding
5. Introduction and Establishment in US
E.
Other
1. Poplar & Willow borer: Cryptorhynchus-hybrid poplars
2. Formosan Termite
32
TREE RESPONSE TO DEFOLIATION
I.
INTRODUCTION
A.
Types of Yield Loss
1.
2.
II.
Direct
a.
Mortality
b.
Radial increment loss
c.
Altered physiological allocation
Indirect
a.
Predisposition to subsequent attack
b.
Susceptibility to environment
c.
Lowered competitive ability
B.
Types of Feeding Behavior
C.
Insect Groups
Physiology of Defoliation
A.
Factors Affecting Response
1.
Hardwoods vs. conifers
2.
Early season vs. late season
3.
Environmental effects
4.
Degrees of and repetitive defoliation
5.
Tissue location and age
B.
Reflushing
C.
The Role of Stress
1.
Effect on plant resistance
2.
Effect on tolerance
33
3.
D.
Changes in host suitability for subsequent attack
1.
Short vs. long-term effects
2.
Types of changes
3.
III.
Population level effects
a.
Allelochemicals
b.
Nutrients
c.
Storage patterns
d.
Phenology
Effects on different feeding guilds
a.
Folivores
b.
Root pathogens
c.
Bark beetles
Relation of Various Control Options to Life History of Defoliating Insects
A.
Biological
B.
Chemical
C.
Silvicultural
34
HARDWOOD DEFOLIATORS
I.
LEPIDOPTERA
A.
Tussock Moths: Lymantriidae
1.
Gypsy Moth: Lymantria dispar
a.
History, current distribution
b.
Life cycle and recognition
c.
Host range and preferences
d.
Population behavior
1.
Weather
2.
Host Plant Quality
3.
4.
e.
a.
Species Preference
b.
Defensive Chemicals: Tannins
Natural Enemies
a.
Generalist Predators
b.
Specialist Parasitoids
c.
Viruses
Interactions Between Host Plants and Natural Enemies
a.
Predators
b.
Parasites
c.
Virus
d.
Btk
Sampling
1.
Pheromones
2.
Egg Mass Survey
3.
Larval Banding
35
f.
Control
1.
Biological
a.
Major natural enemies
Egg:
Ooencyrtus kuvanae (Encyrtidae)
Larva:
Cotesia melanoscela (Braconidae)
Blepharipa intermedia (Tachinidae)
Compsilura concinnata (Tachindae)
NPHV
Entomophaga maimaiga
Pupa: White footed deer mice
Calosoma sycophanta (Carabidae)
Brachymeria intermedia (Cholcididae)
b.
2.
Microbial control
1.
Btk
2.
NPHV
3.
Entomophaga maimaiga
Chemical
a.
Insecticides - Sevin
b.
Pheromones - "Disparlure," "Gyplure"
c.
Insect growth regulators-Dimilin
3.
Silvicultural
4.
Mechanical
g.
Exclusion
h.
Interagency cooperation
i.
Asian gypsy moth: Introductions; Flight in females
36
2.
3.
White-marked tussock moth: Orgyia leucostigma
a.
Host range
b.
Distribution
c.
Biology and recognition
Satin moth: Stilpnotia salicis
Brown-Tail Moth, Nygmia phaeorrhoea
B.
Tent Caterpillars and Webworms
1.
2.
Tent Caterpillars: Lasiocampidae: Malacosoma
a.
Forest Tent Caterpillar: M. disstria
b.
Eastern Tent Caterpillar: M. americanum
c.
Western Tent Caterpillar: M. californicum
Webworms: Arctiidae
a.
C.
Other Arctiidae
1.
D.
Fall Webworm: Hyphantria cunea
Halisidota - "Tussock Moths"
a.
Hickory TM - H. caryae
b.
Spotted TM - H. maculata
c.
Pale TM - H. tessellaris
d.
Sycamore TM - H. harrisii
Inchworms: Geometridae
1.
Inchworms: Geometridae
2.
Fall Cankerworm: Alsophila pometaria
a. distribution
b. host range
c. biology
3.
Spring Cankerworm: Paleacrita vernata
4.
Bruce Spanworm: Operopthera bruceata
5.
Winter Moth: Operopthera brumata
37
E.
F.
G.
II.
1.
Yellow-necked Caterpillar: Datana ministra
2.
Walnut Caterpillar D. integerrima
Royal Moths: Citheroniidae
1.
Hickory Horned Devil: Citheronia regalis
2.
Orange-Striped Oakworm: Anisota senatoria
3.
Green-Striped Mapleworm: Dryocampa rubicunda
Leaf Roller Moths, Leaf tiers, Leaf Miners
COLEOPTERA
A.
B.
III.
Notodontidae
Leaf Beetles: Chrysomelidae
1.
Elm Leaf Beetle: Pyrrhalta luteola
2.
Cottonwood Leaf Beetle: Chrysomela scripta
3.
Willow Leaf Beetle: Plagiorda versicolora
4.
Leaf miners
Scarabaeidae
ORTHOPTERA and PHASMATODEA
A.
Grasshoppers: Acrididae
1.
B.
Red-Legged Grasshopper: Melanoplus femurrubrum
Walkingsticks: Diapheromera femorata
38
IV.
V.
THYSANOPTERA - THRIPS
A.
Introducted Basswood Thrips: Thrips calcaratus
B.
Pear Thrips: Taeniothrips inconsequens
DIPTERA
A.
Leaf Miner Flies: Agromyzidae
1.
VI.
Holly Leaf Miner: Phytomyza ilicicola
HYMENOPTERA (SAWFLIES)
A.
Leaf Miners: Birch, Elm, Alder
B.
Defoliators: Slug Oak Sawfly: Caliroa quercus coccineae
39
CONIFER DEFOLIATORS
I.
BUDWORMS - LEPIDOPTERA: TORTRICIDAE
A.
Eastern Spruce Budworm: Choristoneura fumiferana
1.
Importance, range
2.
Biology
3.
a.
hosts
b.
life cycle
Natural Enemies
a.
Predators - Birds
b.
Parasites
1.
Eggs - Trichogramma minutum (Chalcidae)
2.
Early laravae - Apanteles fumiferanae (Braconidae)
3.
Late larvae - Actia interrupta (Tachinidae)
Meteorus thrichynotus (Braconidae)
4.
c.
4.
5.
Pupal - Itoplectus conquistidor (Ichueumonidae)
Diseases
1.
NPHV
2.
Protozoa - Nosema
3
Fungi
Population dynamics, sampling
a.
cycles
b.
key factors
c.
sampling
Impact
a.
tree
b.
stand
40
6.
B.
C.
II.
Management
a.
silviculture, risk rating
b.
insecticides, IGR's
c.
microbial control
d.
natural enemy enhancement
e.
pheromones
Western Spruce Budworm: Choristoneura occidentalis
1.
Importance, range
2.
Biology
3.
Control
Jack Pine Budworm: Choristoneura pinus
1.
Biology & Host Range
2.
Damage
3.
Control
TUSSOCK MOTHS - LEPIDOPTERA:LYMANTRIIDAE
A.
Douglas fir tussock moth: Orgyia pseudotsugata
1.
Importance, range
2.
Biology
3.
4.
a.
hosts
b.
life cycle
Population dynamics
a.
cycles
b.
factors
c.
sampling
Impact
a.
tree
b.
stand
41
5.
Management
a.
B.
silviculture, risk rating
i.
corrective measures
ii.
preventative measures
b.
insecticides, IGR's
c.
microbial control
d.
natural enemy enhancement
e.
pheromones
Pine Tussock Moth: Dasychira plagiata
1.
Biology
2.
Host range
3.
Site conditions
III. CASEBEARERS AND BAGWORMS
A.
Casebearers (Lepidoptera: Coleophoridae)
1.
B.
a.
Biology
b.
Damage
c.
Control
Bagworms (Lepidoptera: Psychidae)
1.
IV.
Larch Casebearer, Coleophora laricella
Thyridopteryx ephemeraeformis
CONIFER SAWFLIES - HYMENOPTERA: DIPRIONIDAE, TENTHREDINIDAE
A.
General
B.
Importance of Lake States
C.
Redheaded Pine Sawfly - Neodiprion lecontei
1.
Importance
2.
Host range
42
D.
E.
3.
Biology
4.
Control
European Pine Sawfly - N. sertifer
1.
Host range
2.
Biology
Introduced Pine Sawfly - Diprion similis
1.
Host range
2.
Biology
F.
European Spruce Sawfly - D. hercyniae
G.
Larch Sawfly - Pristophora erichsonii
1.
Host range
2.
Biology
V. REGIONALLY IMPORTANT DEFOLIATORS
A.
Needleminers (Lepidoptera: saturniidae)
1.
B.
Silkworm Moths (Lepidoptera: Saturniidae)
1.
C.
Western Hemlock Looper - Lambdina fiscellaria
Sulfur Butterflies - Lepidoptera: Pieridae
1.
E.
Pandora Moth - Coloradia pandora
Inchworms (Lepidoptera: Geometridae)
1.
D.
Lodgepole needleminer - Coleotechnites milleri
Pine Butterfly (Neophasia menapia)
Scarab Beetles (Coleoptera: Scarabaeidae)
1.
Foliar damage
a.
Pine chafer - Anomala oblivia
43
FLUID-FEEDING INSECTS
I.
II.
INTRODUCTION
A.
Nature of Feeding
B.
Nature of Damage
1.
Nutrient loss
2.
Pathogen transmission
3.
Phytotoxins
HEMIPTERA: HOMOPTERA
A.
Adelgids and Aphids
1.
Balsam Woolly Adelgid--Adelges picea
a.
Introduced pest
b.
Host range
c.
Geographic range
d.
Damage
e.
Survey and detection
f.
Control
2. Hemlock Wooly Adelgid, Adelges tsugae
a. Host range
b. Impact
3.
4.
Cooley Spruce Gall Aphid - Adelges cooleyi
a.
complex life cycle, alternate hosts
b.
importance
Pine Bark Aphid - Pineus strobi
a.
5.
Introduced
Gall Aphids
44
B.
C.
SCALE INSECTS
1.
General
2.
Pine Tortoise Scale - Toumeyella parvicornis
3.
Red-pine scale - Matsucoccus resinosae
4.
Beech scale - Cryptococcus fagi
a.
Infection court for Nectria coccinea (Fungus)
b.
Association with lichens
SPITTLEBUGS
1.
Biology
2.
Saratoga Spittlebugs - Aphrophora saratogensis
3.
a.
Biology
b.
Damage
c.
Control
Pine Spittlebug - Aphrophora parallela
a.
b.
Hosts
1.
Preferred - Scotch
2.
Most pines, also Spruce, Fir, Larch, Hemlock
3.
All ages and sizes
Infection counts for Diplodia pini fungus
1.
c.
D.
Control
1.
Thinning: increases vigor
2.
Entomophaga fungus
LEAFHOPPERS: Cicadellidae
1.
White-banded elm leafhopper - Scaphoideus luteolus
a.
E.
Flagging
Elm yellows: Phytoplasma
CICADAS
1.
Ovipositional damage
45
III.
MITES: CLASS ACARINA
A.
Spruce Spider Mite, Oligonychus ununguis
1.
Symptoms
2.
Occurrence following pyrethroid application
IV. OTHER FORMS OF FLUID-FEEDING BY ARTHROPODS
A.
Predation: Hemiptera (true bugs), spiders
B.
Ectoparasitism: Ticks, mites, biting flies
C.
Nectar: Adult moths, bees, flies, etc.
46
SHOOT AND TIP INSECTS
I.
II.
INTRODUCTION
A.
Types of Damage
B.
Interactions with other groups
WEEVILS (COLEOPTERA: CURCULIONIDAE)
A.
White Pine Weevil, Pissodes strobi
1.
Geographic range: Transcontinental
a.
2.
3.
4.
Host range
a.
Northeast and Lake states: White Pine
b.
Pacific Northwest: Sitka Spruce
c.
All pines
Biology
a.
Adult feeding and oviposition
b.
Larval girdling of shoots
Damage
a.
5.
Wisconsin hazard zones
Deformed terminals
Control
a.
Prevention
1.
b.
c.
B.
Hardwood cover
Treatment
1.
Pruning
2.
Removal
Resistance Breeding
Other Weevils
47
III.
SHOOT MOTHS
A.
European Pine Shoot Moth, Rhyacionia buoliana
1.
Geographic range
a.
Northeastern U.S. and Southern Canada
b.
Lake States, S.E. Wisconsin
c.
Pacific Northwest
2.
Host range: All pines, especially Red Pine, <15' tall
3.
Life Cycle
a.
Adults emerge in spring; oviposition at base of buds, needles fascicles,
twig tips.
b.
B.
C.
Larvae
1.
Early Larvae spin webbing; mine needles
2.
Mid Larvae (mid summer): Bore into new buds & overwinter
3.
Late Larvae (spring): Bore into new buds: most damaging phase.
4.
Temperature: Cannot tolerate < -29% C.
Red Pine Shoot Moth, Dioryctria resinosella
1.
Central Wisconsin
2.
30-40 year old trees
3.
Shoots and Cones
Nantucket Pine Tip Moth, Rhyacionia frustrana
1.
Geographic distribution: Eastern, Central and Southern States
2.
Host range: All pines except longleaf and white
3.
a.
South: Loblolly, Shortleaf
b.
Mid-Atlantic: Pitch, Virginia, Scotch
c.
Central: Red
Plantation problems
D.
Zimmerman pine moth, Dioryctria zimmermani
E.
White-Pine Shoot Borer, Eucosma gloriola
48
F.
IV.
1.
Geographic Range: Northeastern and Lake States
2.
Host Range: White, Jack, Red, Scotch Pines
Western Pine Shoot Borer, Eucosma sonomana
BARK BEETLES (Curculionidae: Scolytinae) (Formerly Scolytidae)
A.
B.
Pine Shoot Beetle Tomicus piniperda
1.
Native and North American distribution
2.
Life history
3.
Impact
4.
Management
Pitch Canker
1.
Fusarium subglutinans f. sp. pini, vectored by Pityophthorus twig beetles.
2.
Monterey pine in California.
49
CONE AND SEED INSECTS
I.
CONE AND SEED INSECTS
A.
B.
Introduction
1.
Importance
2.
Ecology
Cone Moths (Lepidoptera)
1.
Red Pine Coneworm, Dioryctria disclusa
2.
Fir Coneworm, Dioryctria abietivorella
a.
C.
D.
True firs, Douglas fir
3.
Spruce Coneworm, Dioryctria reniculelloides
4.
Southern Pine Coneworm
Seed Moths
1.
Spruce Seed Moth, Laspeyresia youngana
2.
Filbertworm
3.
Valentinia
Cone Beetles (Coleoptera)
1.
Bark Beetles (Curculionidae: Scolytinae) (Formerly Scolytidae)
a.
Red Pine Cone Beetle, Conophthorus resinosae
1.
Importance
2.
Hosts
3.
Life cycle
b.
c.
2.
White Pine Cone Beetle, Conophthorus coniperda
1.
Importance
2.
Hosts
3.
Life cycle
Western species of Conophthorus
Weevils (Curculionidae): Curculio
50
E.
F.
a.
Pecan Weevil
b.
Chestnut Weevil
Cone Flies (Diptera)
1.
Spruce Cone Maggot
2.
Walnut Husk Fly
Bugs ( Heteroptera: Hemiptera)
1.
Seed bugs Coreidae - Platycoreia luridans
2.
Pentatomidae - Shieldbacked Pine Seed Bug, Tetyra bipunctata (north),
T. corculus (south)
G.
Gall Wasps (Hymenoptera: Cynipidae)
GALL INSECTS
I.
Major Groups
A.
Aphids
B.
Midges: Cecidomyidae
1.
Balsam Gall Midge
2.
Protection by fungal endophytes
C.
Sawflies - Deciduous: Tenthredinidae
D.
Gall Wasps: Cynipidae
E.
Zimmerman Pine Moth
\
51
APPENDIX 1: INSECT POPULATION SAMPLING TECHNIQUES
I.
Absolute Population Estimates -- expressed as a density (numbers) per unit area
A. Absolute population estimates using marking techniques
1. Methods of marking animals
a. paints and dye solutions
b. labels
c. fluorescent substances (powder)
d. mutilation
e. marking internally by injection
f.
marking by feeding with dyes
g. genes, mutant and normal
h. radioactive isotopes
2. Capture-recapture methods: Lincoln Index
B. Absolute population estimates by sampling a unit of habitat - air, plants,
1. Sampling from the air
a. exposed cone type of suction trap
b. enclosed cone types of suction trap
c. rotary and other traps
2. Sampling from plants
a. direct counting
b. the separation of exposed small animals from the foliage on which they are
living
(1) knockdown by chemicals, jarring and heat
(2) brushing
(3) washing
(4) imprinting
c. the expulsion of insects from trees or shrubs
(1) jarring or beating
(2) chemical knockdown
d. the extraction of insects from herbage and debris
(1) suction apparatus
(2) cylinder or covering method
(3) tents for sampling strongly phototactic insects
(4) extraction by heat and drying
e. methods for insects (eggs) in plant tissues
(1) dissection
(2) bleaching and/or selective staining
(3) X-rays
C. Absolute population estimates by sampling a unit of habitat - soil and litter
1. Mechanical methods of extraction
a. dry sieving
b. soil washing (or wet sieving)
c. soil washing and flotation
d. flotation
e. separation of plant and animal matter by differential wetting
f.
centrifugation
52
g.
h.
a.
D.
II.
III.
sedimentation
elutriation2. Behavioral or dynamic methods
dry extractors
(1) large Berlese funnel
(2) horizontal extractor
(3) multiple canister extractor
(4) Kempson bowl extractor
b. wet extractors
(1) Baermann funnel
(2) hot water extractors
(3) sand extractors
(4) cold water extractor
c. chemical extraction
d. electrical extraction
Absolute population estimates by sampling a unit of habitat -freshwater habitats
Relative Methods of Population Estimation -- the population is measured in unknown
units -- allow only comparisons in space and time; especially useful in extensive work
A. Catch per unit effort
1. Visual observation
2. Flushing
3. Collecting with net or similar device
B. Trapping
1. Interception traps
a. air - flight traps
b. water - aquatic traps
c. land - pitfall and other traps
2. Flight traps combining interception and attraction
a. sticky traps
b. water traps
3. Light and other visual traps
a. shelter traps
b. trap host plants
c. use of vertebrate hosts or substitutes as bait
(1) moving baits
(2) stationary baits
d. bait traps utilizing non-living materials
(1) carbon dioxide and "scents" of animals
(2) carrion and dung
(3) fruits and other attractants
e. sound traps
Estimates based on products and effects of insects (population indices). The insects
themselves are not counted, but their products (e.g., frass, webs, exuviae, nests, etc.,) or
effects (e.g., plant damage) are counted.
A. Products
1. Exuviae
53
2. Frass
3. Other products
B. Effects
1. Economic damage
2. Yield
3. Amount of plant consumed
54
APPENDIX 2: MAJOR CATEGORIES OF INSECTICIDES
I.
PYRETHROIDS
A. Mode of Action - Axon poison
B. Attributes Moderately broad acting
Highly effective; very low doses needed
Moderately persistent in environment
Low mammalian toxicity
High fish toxicity
May release mite outbreaks
C. Examples - Pydrin
D. Comments Expensive
Derived from Chrysanthemum
II.
ORGANOPHOSPHATES
A. Mode of Action - Acetylcholinesterase inhibitor
B. Attributes Very broad acting
Less persistent in environment
High mammalian toxicity
Some miticidal activity
C. Examples Malathion, orthene, lorsban, cygon
D. Comments Inexpensive; commonly used
III.
CARBAMATES
A. Mode of Action - Acetylcholinesterase inhibitor
B. Attributes Moderately broad acting
Less persistent in environment
Moderate mammalian toxicity (sometimes high)
Sometimes high bee (and earthworm) toxicity
C. Examples Carbaryl "Sevin," Temik, Vydate
D. Comments Sevin is widely used
IV.
NEONICOTINOIDS
A. Mode of Action - Postsynaptic inhibitor
B. Attributes Moderately broad acting
Less persistent in environment
Usually low mammalian toxicity
Widely used against sucking insects, boring insects because of
systemic activity
C. Examples Imidacloprid very widely used
D. Comments Derived from nicotine
V.
MICROBIAL PESTICIDES
A. Bacillus thuringiensis
1. Different subspecies used on Lepidoptera, Coleoptera, Diptera
2. Attributes
Moderately broad acting
Less persistent in environment
55
B.
No known mammalian toxicity
Widely used
3.
Examples Dipel
Viruses
1. Attributes
Species specific
Less persistent in environment
No known mammalian toxicity
Expensive, sometimes slow acting
3.
Examples Gypchek
VI. INSECT GROWTH REGULATORS
VII.
ORGANOCHLORINES
A. Mode of Action - Axon poison
B. Attributes Very broad acting
Highly persistent in environment
Longterm control
Environmental Contamination; Biomagnification
Bird egg shell thinning
Variable toxicity, carcinogenicity & mutagenicity
May releases mite outbreaks
C. Examples DDT, lindane, chlordane
D. Comments Most are banned for most uses
INSECTICIDE TERMINOLOGY
Systemic LD50 LT50
-
Will be taken up by plant
The dose required to kill 50% of the population; the lower the LD50, the higher the
toxicity
The time required to kill 50% of the population
MECHANISMS OF INSECTICIDE RESISTANCE
Reduced penetration
Excretion
Sequestration
Detoxification
Target Site Insensitivity
For resistance management, see recommendations by IRAC
56