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The Bark Beetles, Fuels, and Fire Bibliography Quinney Natural Resources Research Library, S.J.
and Jessie E.
1-1-2005
Root Diseases in Coniferous Forests of the Inland
West : Potential Implications of Fuels Treatments
Raini C. Rippy
Jane E. Stewart
Paul J. Zambino
Ned B. Klopfenstein
Joanne M. Tirocke
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Recommended Citation
Rippy, R., Stewart, J., Zambino, P., Klopfenstein, N., Tirocke, J., Kim, M-S. and Thies, W. (2005). Root diseases in coniferous forests
of the inland west : potential implications of fuels treatments. USDA Forest Service, Rocky Mountain Research Station, General
Technical Report RMRS-GTR-141, 26 pp.
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The Authors _________________________________________
Raini C. Rippy is a Biological Science Technician with the USDA Forest Service, Rocky
Mountain Research Station at the Forestry Sciences Laboratory in Moscow, ID, and is pursuing
a doctorate in plant pathology at Washington State University. She completed a B.S. degree in
forest resources and an M.S. degree in forest products at the University of Idaho.
Jane E. Stewart is a Biological Science Technician with the USDA Forest Service, Rocky Mountain
Research Station at the Forestry Sciences Laboratory in Moscow, ID.
Paul J. Zambino is a Research Plant Pathologist with the USDA Forest Service, Rocky Mountain
Research Station at the Forestry Sciences Laboratory in Moscow, ID.
Ned B. Klopfenstein is a Research Plant Pathologist with the USDA Forest Service, Rocky
Mountain Research Station at the Forestry Sciences Laboratory in Moscow, ID.
Joanne M. Tirocke is a Biological Science Technician with the USDA Forest Service, Rocky
Mountain Research Station at the Forestry Sciences Laboratory in Moscow, ID.
Mee-Sook Kim is a Plant Pathologist with the USDA Forest Service, Rocky Mountain Research
Station at the Forestry Sciences Laboratory in Moscow, ID.
Walter G. Thies is a Research Plant Pathologist with the USDA Forest Service, Pacific
Northwest Research Station in Corvallis, OR.
Abstract ____________________________________________
Rippy, Raini C.; Stewart, Jane E.; Zambino, Paul J.; Klopfenstein, Ned B.; Tirocke, Joanne M.; Kim, Mee-Sook;
Thies, Walter G. 2005. Root diseases in coniferous forests of the Inland West: potential implications
of fuels treatments. Gen. Tech. Rep. RMRS-GTR-141. Fort Collins, CO: U.S. Department of Agriculture,
Forest Service, Rocky Mountain Research Station. 32 p.
After nearly 100 years of fire exclusion, introduced pests, and selective harvesting, a change in forest
composition has occurred in many Inland West forests of North America. This change in forest structure
has frequently been accompanied by increases in root diseases and/or an unprecedented buildup of fuels.
Consequently, many forest managers are implementing plans for fuels treatments to lower the risk of
severe wildfires.
Impacts on root disease should be considered before selecting appropriate fuels treatments. Complex
interactions exist among conifer root diseases, fuels treatments, forest structure, species composition,
stand history, and other environmental factors. As forest managers prescribe fuels treatments, their
success in lowering the risk of severe wildfire will depend in part on the impacts of these treatments on root
disease. Root diseases are one of many factors to be considered when developing plans for fuels
treatments. Choices must be made on a site-by-site basis, with knowledge of the diseases that are present.
This paper provides examples of how fuels treatments may increase or reduce specific diseases and
demonstrates their importance as considerations in the fuels management planning process. Several root
diseases prevalent within Inland West of North America are addressed: Armillaria root disease, annosus
root disease, laminated root rot, black stain root disease, Schweinitzii root and butt rot, Tomentosus root
disease, Rhizina root rot, and stringy butt rot. For each disease, general information is provided on disease
identification, management options, and potential effects of fuels treatments. However, many long-term
studies are needed to assess effects of specific interactions among fuels treatments, root diseases, and
host trees.
Key words: wildfire, forest planning, forest structure, mechanical treatments, prescribed fire,
coniferous hosts, pathogens, disease management
Foreword ______________________
This document is part of the Fuels Planning: Science
Synthesis and Integration Project, a pilot project initiated
by the USDA Forest Service to respond to the need for
tools and information useful for planning site-specific fuel
(vegetation) treatment projects. The information ad-
dresses fuel and forest conditions of the dry inland
forests of the Western United States: those dominated by
ponderosa pine, Douglas-fir, dry grand fir/white fir, and
dry lodgepole pine potential vegetation types. Informa-
tion was developed primarily for application at the stand
level and is intended to be useful within this forest type
regardless of ownership. Portions of the information also
will be directly applicable to the pinyon pine/juniper
potential vegetation types. Many of the concepts and
tools developed by the project may be useful for planning
fuel projects in other forests types. In particular, many of
the social science findings would have direct applicability
to fuel planning activities for forests throughout the United
States. As is the case in the use of all models and
information developed for specific purposes, our tools
should be used with a full understanding of their limita-
tions and applicability.
Root Diseases in Coniferous Forests of the Inland
West: Potential Implications of Fuels Treatments
The science team, although organized functionally,
worked hard at integrating the approaches, analyses,
and tools. It is the collective effort of the team members
that provides the depth and understanding of the work.
The science team leadership included the following USDA
Forest Service personnel:
•Deputy Science Team Leader Sarah McCaffrey,
North Central Research Station
•Forest structure and fire behavior: Dave Peterson
and Morris Johnson, Pacific Northwest Research
Station
•Environmental consequences: Elaine Kennedy-
Sutherland and Anne Black, Rocky Mountain Re-
search Station
•Economic uses of materials: Jamie Barbour and
Roger Fight, Pacific Northwest Research Station
•Public attitudes and beliefs: Pamela Jakes and
Susan Barro, North Central Research Station
•Technology transfer: John Szymoniak, Pacific
Southwest Research Station
This project would not have been possible were it not
for the vision and financial support of the USDA Forest
Service Washington Office Fire and Aviation Manage-
ment individuals, Janet Anderson Tyler and Leslie
Sekavec.
Russell T. Graham, Science Team Leader
USDA Forest Service, Rocky Mountain
Research Station
Acknowledgments _________________________
This paper was supported by the USDA Forest Service, Fuels Planning:
Science Synthesis and Integration Project, Rocky Mountain Research Station
RWU-4552 (Microbial Processes as Ecosystem Regulators in Western For-
ests), and the Sustainable Forestry component of Agenda 2020, a joint effort
of the USDA Forest Service Research and Development and the American
Forest and Paper Association. The authors thank Drs. Deborah S. Page-
Dumroese and Russell T. Graham for their support on this project. We thank
Dr. Gregory M. Filip, Dr. John W. Schwandt, Dr. Susan K. Hagle, Dr. Jack D.
Rogers, Ms. Kris L. N. Fields, Mr. Brennan A. Ferguson, Ms. Jonalea R. Tonn,
Ms. Ann Abbott, and Mr. Bob Atwood for their comments on an earlier version
of this manuscript. We also thank Dr. Jack D. Rogers, Dr. John W. Schwandt,
and Mr. John W. Hanna for their contributed photographs. In addition, we
greatly appreciate the helpful reviews by three anonymous reviewers.
Contents ___________________________________
Page
Introduction .................................................................................................. 1
Armillaria Root Disease................................................................................ 6
Disease Symptoms and Signs ......................................................... 6
Disease Management .................................................................... 10
Fuels Treatment Considerations .................................................... 10
Annosus Root Disease............................................................................... 11
Disease Symptoms and Signs ....................................................... 11
Disease Management .................................................................... 13
Fuels Treatment Considerations .................................................... 14
Laminated Root Rot ................................................................................... 15
Disease Symptoms and Signs ....................................................... 15
Disease Management .................................................................... 15
Fuels Treatment Considerations .................................................... 17
Black Stain Root Disease........................................................................... 18
Disease Symptoms and Signs ....................................................... 18
Disease Management .................................................................... 20
Fuels Treatment Considerations .................................................... 20
Schweinitzii Root and Butt Rot ................................................................... 21
Disease Symptoms and Signs ....................................................... 21
Disease Management .................................................................... 21
Fuels Treatment Considerations .................................................... 21
Other Root Diseases of Consideration in the Inland West ......................... 23
Tomentosus Root Disease ............................................................. 23
Rhizina Root Rot ............................................................................ 23
Stringy Butt Rot .............................................................................. 23
Disease Management and Fuels Considerations .......................... 24
Summary of Fuels Management Considerations ....................................... 25
References ................................................................................................. 26
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1
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Introduction ____________________
The effects of nearly 100 years of fire exclusion, introduced
pests, and selective timber harvesting have caused dramatic
changes in tree species composition across much of the West-
ern landscape. These changes have contributed to an unprec-
edented buildup of fuels and increased incidence of root
disease (Tiedemann and others 2000). Heavy accumulations of
fuels and ample fuel ladders have put stands at risk for
uncharacteristically severe wildfires. Consequently, many for-
est managers are implementing plans for fuels treatments to
lower the risk of fire. An understanding of how these fuels
treatments might affect the many complex interactions that
exist within forest ecosystems is critical because even subtle
disturbances can trigger associated responses from many co-
existing organisms (Borchers and Perry 1990).
Historically, root diseases have played an important and
beneficial role in ecological processes of succession, decom-
position, and fire in forests with moderate- to high-intensity
fire regimes (Hessburg and others 1994; Hansen and Goheen
2000). As with other tree pathogens, native root pathogens are
thought to have coevolved with their hosts and are a natural,
and even necessary, part of forest ecosystems. In most situa-
tions, native root diseases do not cause irreplaceable loss of
entire stands over large areas nor threaten the existence of any
host species. However, shifts in stand composition and other
natural and human-caused disturbances have frequently re-
sulted in increased damage from root-rot diseases (Edmonds
and others 2000). In some situations, the change in root rot
dynamics may influence forest growth and succession for
centuries. Thus, careful consideration of future root disease
impacts is critical before implementing any management ac-
tivities, including fuels treatments.
Root Diseases in Coniferous
Forests of the Inland West:
Potential Implications of Fuels
Treatments
Raini C. Rippy
Jane E. Stewart
Paul J. Zambino
Ned B. Klopfenstein
Joanne M. Tirocke
Mee-Sook Kim
Walter G. Thies
Whether specific effects of root diseases are considered to be
beneficial or detrimental depends on social values and man-
agement objectives for the site. For some sites, impacts by root
disease in a stand may be viewed as undesirable because of
potential increases in fuels buildup, reduced timber volume,
damage to facilities from wind-thrown trees, and potential
personal injury. At other sites, with different management
objectives, loss of trees as a result of root disease may be
acceptable or even desirable because of benefits attributed to
disease-created openings, scattered mortality (lower density
stand), and increased volume of standing and downed large
woody debris. Such disease-associated changes may provide
forage for large game animals, habitat for small animals and
cavity-nesting birds, sources for specialty forest products,
increased diversity in the plant community, and other changes
in forest structure that reduce the likelihood of catastrophic
wildfire (Thies 1999; Filip 1999; Thies and Goheen 2002).
Fuels treatments range widely in approach and application,
but generally involve mechanical treatments and/or prescribed
fire (Fitzgerald 2002; Graham and others 2004; Peterson and
others 2005). Mechanical treatments, such as thinning, can
remove fuels as usable forest products, which can include
biomass energy. Other types of treatments may involve cutting
trees, reducing the size of fuels, eliminating fuels ladders, and
methods that place fuels on the forest floor to accelerate
decomposition, such as mowing (Fitzgerald 2002). Prescribed
fire is also used to reduce surface fuels and ladder fuels;
however, the effects of prescribed fire on resulting stand
structure are less predictable than mechanical treatments. As
forest managers implement plans for fuels treatments, their
success in lowering risk of severe wildfire will depend in part
on the impacts of management activities on levels of root
disease that contribute to subsequent fuels accumulation. The
2USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
type of fuels treatment can often be selected to match the
overall objectives for a stand, while addressing root rot dynam-
ics and possible detrimental effects on a site.
Fuels management practices can influence root disease in
complex ways that should be considered at the ecosystem
level. Tree thinning can generate stumps and residual root
systems that are susceptible to colonization by root-rot patho-
gens. Stumps, roots, and slash can serve as nutrient substrates
that increase the overall growth and distribution of root rot
fungi. Thinning-associated wounds on retained trees can pro-
vide opportunities for root disease fungi to cause new infec-
tions. When seral tree species—for example, pine and larch—
are removed in thinning operations, the resulting stand may
consist of a higher proportion of tree species that are more
susceptible to root rots, such as grand fir or Douglas-fir. Fuels
treatments can cause physiological stress to remaining trees,
increasing the likelihood that root rot problems will develop.
Although the effects of slash burning may vary, superficial
burning appears to have little impact on the distribution of
pathogens, such as Armillaria species, Phellinus weirii, or
Heterobasidion annosum, which frequently occupy protected
areas within structural roots (Hadfield and others 1986). How-
ever, some measures that limit substrate availability for root
pathogens may reduce inoculum and possibly reduce disease
severity. Thus, care must be taken to select appropriate fuels
treatments that will achieve the objective of reducing current
and future fuels levels while avoiding undesirable increases in
root disease. Evaluating the presence and distribution of root
diseases is an essential step in selecting appropriate fuels
treatments. Surveys for root disease include observing signs
and symptoms on living trees, and also examining snags,
stumps, and overturned trees (Thies 1995).
Although fuels reduction has recently emerged as a national
issue, there are few research studies specifically addressing
fuels-treatment effects on root diseases. For this reason, much
of the information applied to fuels treatments in this paper is
based on general principles of plant pathology, knowledge of
pathogen biology, and/or derived from studies conducted for
other purposes. Because the consequences of root disease
resulting from fuels treatments may not be immediately appar-
ent but may continue for decades after treatments, there is an
increased need to conduct and interpret long-term studies that
examine effects of specific fuels treatment on root disease
interactions. Such long-term studies would provide more pre-
cise information for understanding these complex relation-
ships.
The objectives of this paper are to (1) provide insights for
consideration regarding potential effects of fuels treatments on
root diseases, (2) identify specific needs for research pertain-
ing to the effects of fuels treatments on root diseases, and (3)
aid field personnel in identification of the primary root dis-
eases. This discussion focuses on the dry, mixed conifer forests
between the crest of the Cascade Mountains and the crest of the
Rocky Mountains in North America, hereafter referred to as
the Inland West. In forests of the Inland West, special attention
is warranted for several root diseases, such as Armillaria root
rot (caused by Armillaria species), Annosus root rot (caused by
Heterobasidion species), laminated root rot (caused by Phellinus
weirii), black stain root disease (caused by Leptographium
wageneri), and Schweinitzii root rot (caused by Phaeolus
schweinitzii), and other diseases including Tomentosus root
disease (caused by Inonotus tomentosus), Rhizina root rot
(caused by Rhizina undulata), and stringy butt rot (caused by
Perenniporia subacida). Rudimentary information is pro-
vided in this paper for identification of these diseases; how-
ever, more detailed and definitive field guides with a greater
representation of pathogens are available (Partridge and Miller
1974; Partridge and others 1978; Hadfield and others 1986;
Scharpf 1993; Allen and others 1996; Filip 1999; Hagle and
others 2003). Figures illustrate signs and symptoms of the
more common root diseases. We included factors to be consid-
ered in the effects of potential fuels treatments on each root
disease and identify gaps in research information. Three tables
located on the next 3 pages contain general supporting infor-
mation: table 1 lists common symptoms and signs of each root
disease; table 2 lists common coniferous hosts of each root
disease; and table 3 lists common and scientific names of
pathogens, trees, and insect species.
3
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Table 1—Symptoms and signs associated with common root diseases of conifers of the Inland West.
Root disease Symptoms Signs
(causal fungus) (the most diagnostic are in bold text) (the most diagnostic are in bold text)
Armillaria root disease Reduced tree growth Mycelial fans within tightly attached bark
(
Armillaria ostoyae
)Crown thinning and yellowing Rhizomorphs
Smaller, stress cones Fruiting bodies:
Resinosis (sap on butt/roots) Honey mushrooms in late summer or fall
Wood decay/rot: spongy, yellow, stringy
Annosus root disease Reduced tree growth Ectotrophic mycelium on roots
(
Heterobasidion annosum
)Reduced growth of lateral branches Fruiting bodies:
Crown thinning and yellowing Perennial conks:
Smaller, stress cones dark brown (top)
Red staining in the heartwood cream colored, porous (below)
Wood decay: First-year conks
Spongy, stringy, white with black flecks buff-colored when immature
Laminated decay
Black stain root disease Reduced tree growth Dark purple-to-black hyphae:
(
Leptographium wageneri
)Crown thinning Stain in recent sapwood and cambium
Stress cones Lengthwise stain in lower bole
Rapid crown wilting
Laminated root rot Reduced tree growth Ectotrophic mycelium
(
Phellinus weirii
)Crown thinning and yellowing Reddish-brown setal hyphae between
Laminated decay in roots/lower bole layers of decayed wood
Layers of decay contain small pits Fruiting bodies:
Red-brown staining flat, cream-brown
Rhizina root disease Only in recently burned area Masses of white to yellow mycelia form
(
Rhizina undulata
)Dead seedlings mats around roots
Sparse foliage Fruiting bodies:
Discolored needles Disklike, with multiple rootlike
Stem resinosis structures scattered across underside
Groups form along or over infected roots
Brown to black with age
Schweinitzii root and butt rot Reduced tree growth Fruiting bodies:
(Red-brown butt rot) Crown thinning and yellowing On ground: circular or irregularly lobed
(
Phaeolus schweinitzii
)Gall-swelling of small roots On trees: shelf-like
Pronounced butt swelling Fresh: Upper surface velvety, usually
Yellow/red-brown staining in heartwood reddish-brown with a yellow margin
Wood decay: Older: brown, resembling cow dung
Cubical decay, reddish-brown, crumbly
Carpenter ants often occur within decay
Stringy butt rot Wood decay: Yellow mycelial mats between layers of
(
Perenniporia subacida
)
Light brown stain in heartwood decayed wood
Small white pits in decayed wood, Fruiting bodies:
Masses of stringy, white spongy fibers On undersides of downed trees and
with small black flecks branches of living trees
Laminated wood decay Perennial, flat and leathery to crustlike,
cream to light yellow, underside covered in
small circular pores
Tomentosus root rot Resin exuded on infected roots Fruiting bodies:
(
Inonotus tomentosus
) Wood decay: Annual, circular, on roots or ground, often
Red stain, small pockets lined with united with multiple stems.
white fibers Upper surface: velvety, yellow to brown
Lower surface: light brown pores darken
when bruised
4USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Table 2—Common coniferous hosts of root diseases of the Inland West. Consult text for major and minor hosts for each
pathogen species, form, type, and variety.
Coniferous hosts
Hemlocks, cedars,
Root disease Pines Douglas-fir and firs Spruce and larch juniper, and yew
Armillaria root disease Jeffrey pine Douglas-fir Engelmann spruce juniper
lodgepole pine grand fir mountain hemlock
ponderosa pine subalpine fir western hemlock
western white pine white fir western redcedar
whitebark pine
Annosus root disease Jeffrey pine Douglas-fir Engelmann spruce incense cedar
ponderosa pine grand fir juniper
white fir mountain hemlock
other true firs western hemlock
Black stain root disease Jeffrey pine Douglas-fir Can be found on a Can be found on
lodgepole pine variety of spruce hemlock species, but
pinyon pine species, but found found only occasionally
ponderosa pine only occasionally
western white pine
Laminated root rot Can be found on Douglas-fir Engelmann spruce mountain hemlock
pines, but not grand fir western larch Pacific yew
common subalpine fir western hemlock
white fir
Rhizina root rot lodgepole pine Douglas-fir Englemann spruce western hemlock
ponderosa pine grand fir spruce species western redcedar
western white pine western larch
other pine species
Schweinitzii root lodgepole pine Douglas-fir western larch western hemlock
and butt rot ponderosa pine true firs many spruce species western redcedar
western white pine
Stringy butt rot lodgepole pine Douglas-fir Engelmann spruce western hemlock
western white pine true firs western larch western redcedar
white spruce
Tomentosus root rot lodgepole pine Douglas-fir Engelmann spruce
ponderosa pine grand fir white spruce
western white pine white fir
5
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Table 3—Common and scientific names for pathogens, tree species, and insect species.
Root diseases Scientific name of causal fungus
Armillaria root disease
Armillaria ostoyae
Annosus root disease
Heterobasidion annosum
Black stain root disease
Leptographium wageneri
Laminated root rot
Phellinus weirii
Rhizina root rot
Rhizina undulata
Schweinitzii root and butt root (Red-brown butt rot)
Phaeolus schweinitzii
Stringy butt rot (Yellow root rot)
Perenniporia subacida
Tomentosus root rot
Inonotus tomentosus
Trees Scientific name
Aspens, Cottonwoods
Populus spp.
Douglas-fir
Pseudotsuga menziesii
Engelmann spruce
Picea engelmannii
Grand fir
Abies grandis
Incense cedar
Calocedrus decurrens
Jeffrey pine
Pinus jeffreyi
Juniper
Juniperus spp.
Larch
Larix spp.
Lodgepole pine
Pinus contorta
Mountain hemlock
Tsuga mertensiana
Oaks
Quercus spp.
Pacific yew
Taxus brevifolia
Ponderosa pine
Pinus ponderosa
Singleleaf pinyon
Pinus monophylla
Spruce
Picea spp.
Subalpine fir
Abies lasiocarpa
Sugar pine
Pinus lambertiana
Western hemlock
Tsuga heterophylla
Western larch
Larix occidentalis
Western redcedar
Thuja plicata
Western white pine
Pinus monticola
Whitebark pine
Pinus albicaulis
White fir
Abies concolor
Insects Scientific name
Douglas-fir beetle
Dendroctonus pseudotsugae
Douglas-fir pole beetle
Pseudohylesinus nebulosus
Fir engraver beetle
Scolytus ventralis
Mountain pine beetle
Dendroctonus ponderosae
Western pine beetle
Dendroctonus brevicomis
6USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Armillaria Root Disease __________
Armillaria root disease, also known as shoestring root rot, is
caused by a variety of Armillaria species that are widely
distributed throughout the Western United States, Europe,
Asia, Africa, and Australia (Watling and others 1991). In
North America, Armillaria ostoyae (Romagn.) Herink pre-
dominantly infects conifers; whereas other species of Armillaria
primarily infect hardwoods or are primarily saprophytic (Gre-
gory and others 1991). Ecological roles of Armillaria species
include those of primary pathogen, secondary invader, and
saprophyte. As a primary pathogen, Armillaria is capable of
killing vigorous hosts, while as a secondary invader it only
colonizes hosts predisposed to disease. Armillaria ostoyae is
the most widespread and aggressive species of Armillaria on
conifers in forests of the Inland West (Kile and others 1991).
It can survive up to several decades in previously infected
stumps (Goheen and Otrosina 1998). A recent study estimated
that a single pathogenic A. ostoyae individual occupied an area
of 965 ha (2,385 acres) in a mixed-conifer forest of northeast-
ern Oregon and estimated its age to be between 1,900 and 8,560
years old (Ferguson and others 2003). Thus, Armillaria root
disease can be a persistent, long-term influence within forest
ecosystems. Armillaria root disease causes significant mortal-
ity, wood decay, and growth reduction in infected conifers
(Williams and others 1989).
The host range for A. ostoyae includes several western
conifer species in North America; however, susceptibility of
individual tree species may vary depending on site factors and
location (Goheen and Otrosina 1998). Armillaria ostoyae
causes more mortality in forests of the Inland West than in
coastal forests (Goheen and Otrosina 1998), but Armillaria
root disease is a serious problem in both regions (Morrison
1981; Hadfield and others 1986). Highly susceptible hosts
include Douglas-fir, grand fir, and white fir. Other species
damaged or killed by A. ostoyae include ponderosa pine,
Jeffrey pine, lodgepole pine, whitebark pine, western white
pine, Engelmann spruce, subalpine fir, western and mountain
hemlock, western redcedar, and juniper. Some hardwoods
including species of oak, cottonwood, and aspen are damaged
or killed by other species of Armillaria, such as A. mellea
(Morrison 1981; Hadfield and others 1986; Filip 1999; Filip
and Ganio 2004).
Host range, virulence, and other aspects of the ecological
behavior of Armillaria species vary with geographic location.
It can be difficult to predict where Armillaria root disease
might occur (McDonald and others 1987a,b; Kile and others
1991; Mallett and Maynard 1998). On some sites, pines and
other species are tolerant to A. ostoyae, but on other sites
almost all tree species are killed (Goheen and Otrosina 1998).
Locations where trees are stressed—such as sites with drain-
age problems, soil compaction, or those on the outer limits of
geographic range for the host—have increased mortality (Close
1988; Gregory and others 1991; Goheen and Otrosina 1998).
Occurrence of Armillaria root disease appears to be associated
with groups of habitat types called vegetation subseries
(McDonald and others 2000). Pathogenic Armillaria is rare or
absent in hot-dry, cold-dry, and frost-pocket habitat types
(McDonald 1998, 1999). Conifers growing on coarse-textured
soils are typically at greater risk of developing Armillaria root
disease than those growing on fine-textured soils (Mallett and
Maynard 1998).
Armillaria root disease is commonly associated with other
root diseases, and it may be difficult to tell which disease
occurred first or which disease has caused the most damage.
Additionally, Armillaria root disease may predispose conifers
to bark beetle attacks (Partridge and Miller 1972; Lane and
Goheen 1979; Schmitt and others 1984; Schowalter and Filip
1993).
Disease Symptoms and Signs
Most symptoms of Armillaria root disease (reduced height
growth, thinning and yellowing of foliage, and/or abundant
crops of smaller stress cones) are common to many root
diseases. Although not always present, pitch flow just above
the ground is a fairly diagnostic symptom on resinous conifer
species (fig. 1). Signs that are indicative of Armillaria root
disease can be found at the base, roots, and beneath the bark of
an infected tree or stump. Signs include:
•Mycelial mats or fans (white, thick, fan-like shapes that
can be found under the bark) (fig. 2).
•Rhizomorphs (reddish-black, flat, shoestring-like myce-
lial cords that are commonly present on or beneath the
bark of roots and lower stems and in the soil near infected
trees or stumps).
•Presence of honey-colored mushrooms (the fruiting bod-
ies of the pathogen) produced in late summer or fall after
rain (fig. 3) (Morrison 1981; Hadfield and others 1986;
Williams and others 1989).
Fans and rhizomorphs do not always indicate that mortality
has been caused by Armillaria root disease. Pathogenic
Armillaria is diagnosed if mycelial fans are found within bark
that is tightly attached to living wood, especially if resinosis is
present at the root crown. Extensive growth of the fungus
eventually girdles and kills the tree. As mycelial fans age, they
become yellow and begin to disintegrate, but a fan-shaped
imprint is left behind in the wood and bark (Goheen and
Otrosina 1998). Pitch flow is an indication of the living tree’s
response to the pathogenic fungus, and the amount of resin
flow may be indicative of the amount of A. ostoyae-caused
colonization in the roots (Omdal and others 2004). In contrast,
fans of saprophytic species of Armillaria often invade between
loosened bark and wood of dead trees or trees dying from other
causes. Trees invaded by saprophytic Armillaria spp. exhibit
little or no evidence of resinosis or wound callus formation
(Robinson and others 2004).
As the disease progresses, stands affected by Armillaria root
disease display increased mortality, often with uprooted trees
with roots broken at some distance from the bole (fig. 4 through
6). Also, as infected trees decay, the cambial tissues and wood
7
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Figure 1—Abundant resin flow at the base and roots of a
conifer is a common symptom indicating infection by
Armillaria
ostoyae
. (Photo courtesy of R.C. Rippy)
Figure 2—Mycelial fans are present within roots of this grand fir infected
with
Armillaria
ostoyae
. (Photo courtesy of J.W. Hanna)
8USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Figure 3—Fruiting bodies of
Armillaria
species.
Armillaria ostoyae
commonly fruits in large clusters; however, mushrooms of other
Armillaria
species also grow in clusters. (Photo courtesy of R.C. Rippy)
Figure 4—In this
Armillaria
-caused mortality center, mushrooms
from a species of
Armillaria
can be seen in the foreground.
(Photo courtesy of J.W. Hanna)
9
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Figure 5—
Armillaria
-caused mortality
centers as they appear in mixed conifer
stands. (Photos courtesy of J.D. Rogers)
Figure 6—An uprooted ponderosa pine showing
decayed roots that have broken at some distance
from the bole. These root wad characteristics, with
other signs and symptoms, indicate mortality from
Armillaria root disease. (Photo courtesy of R.C.
Rippy)
10 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
become yellowish, moist, stringy, and sometimes have a
pungent odor (Goheen and Otrosina 1998). In the later stages
of decay, wood becomes spongy, light yellow, and may be
marked with numerous black lines, called “zone lines”
(Morrison and others 1992). Decayed wood becomes hard and
fibrous as it dries, as can be seen in wind-blown trees. Mush-
rooms produced by Armillaria species are often called honey
mushrooms because of the honey-colored caps produced by
some species (fig. 3). The mushrooms have white spores, gills
attached to the stem, minute peg-like hairs at the center of the
upper cap, and a cottony ring around the upper stem (stipe).
These mushrooms are often found growing in clusters near the
bases of infected trees or stumps (fig. 4). Although, mush-
rooms are typically produced during moist conditions in the
late summer and fall, they are sometimes difficult to find, and
they are not produced every year. Other species of Armillaria
produce similar mushrooms, which can make field identifica-
tion unreliable.
Disease Management
Armillaria root disease is difficult to control because the
disease inoculum (mycelium and rhizomorphs that have colo-
nized woody substrates) is nearly impossible to eliminate from
a site. In timber-growing areas, management decisions should
consider site characteristics (habitat type, soil type, and asso-
ciated conditions), stand structure, host species composition,
stand management history, and Armillaria species present to
minimize impacts of this disease (Morrison 1981; Hadfield
and others 1986; Williams and others 1989; Fox 2000). Im-
pacts of Armillaria root disease can be reduced by:
•Favoring more resistant/tolerant tree species.
•Maintaining tree species diversity.
•Reforesting stands with locally adapted species suitable
to the site.
•Promoting tree vigor by minimizing stress and avoiding
wounds.
•Reducing inoculum sources through the uprooting of
stumps and removal of woody debris (Williams and
others 1989; Hagle and Shaw 1991; Roth and others
2000).
These management considerations are especially critical for
sites with high levels of inoculum and/or conditions favorable
for disease development (Hagle and Shaw 1991). Thinning has
also been examined as a possible management option for
reducing Armillaria root disease within a stand. Some studies,
outside of the Inland West, in the Cascade Range of western
Oregon and Washington have shown that precommercial thin-
ning either decreases or does not impact mortality caused by
Armillaria root disease (Filip and others 1989; Filip and Ganio
2004). Precommercial thinning may improve tree vigor by
decreasing tree stress, thereby reducing infection. In contrast,
thinning increased the mortality caused by the disease in
several Inland West locations, such as northern Idaho and parts
of Canada (Cruickshank and others 1997). Stumps that remain
after thinning treatments are a potential inoculum source that
could increase new infections (Cruickshank and others 1997).
Differences in the results from these studies highlight that site
location and plant association can potentially affect outcomes
of management choices, and suggest that these factors be
considered in planning fuels treatments. More research infor-
mation about Armillaria root disease has been compiled in
Shaw and Kile (1991).
Fuels Treatment Considerations
Because Armillaria species can reside deep in the soil and
within large woody roots, superficial slash burning or pre-
scribed fire is unlikely to eliminate the pathogen, although
these treatments may reduce infection potential indirectly by
removing highly susceptible host species that are readily killed
by fire (Hadfield and others 1986; Filip and Yang-Erve 1997;
Ferguson and others 2003). Mortality caused by Armillaria
increases fuel loads (Fields 2003). Prescribed fire may be used
to reduce fuel loads, which may also reduce inoculum levels
above ground by eliminating nutritional substrates for growth
of Armillaria. However, fire may have minimal direct effects
on the pathogen below ground (Filip and Yang-Erve 1997).
Yet, fire may have indirect effects on Armillaria inoculum by
increasing populations of antagonistic soil fungi such as Tri-
choderma (Reaves and others 1990). Studies have demon-
strated that Armillaria species survive intense fires (Ferguson
and others 2003) and can readily colonize roots of trees killed
by fire (Blodgett and Lundquist 2004). Fire may increase or
decrease the susceptibility of a stand through its influence on
species composition, stand structure, and tree vigor. Thinning
treatments that favor climax or late-successional host trees will
likely aggravate Armillaria root disease. However, thinning
that favors early seral species may reduce root disease over the
long term. Also, greater spacing can improve tree vigor of seral
species, which may increase tolerance to Armillaria root dis-
ease.
Recently, a Web-based tool (Armillaria Response Tool:
ART) has been developed that can estimate risk of Armillaria
root disease caused by A. ostoyae in dry forests of the Western
United States (McDonald and others, in press). This tool uses
habitat types to identify sites where A. ostoyae may occur, and
the tool indicates how some fire reduction activities (fuels
management) may exacerbate Armillaria disease within high-
risk stands.
11
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Annosus Root Disease ___________
Annosus root disease, caused by Heterobasidion annosum
(Fr.) Bref., is found in many temperate coniferous forests
around the world. It is an endemic pathogen that is common
and widely distributed in North America. Although this dis-
ease has existed for millennia within coniferous forest ecosys-
tems, harvesting practices have increased its incidence and
impacts, especially in Western and Southeastern North America.
In many forest stands, annosus root disease has reached epi-
demic levels (Otrosina and Ferrell 1995).
Within the Inland West, annosus root disease is of most
concern on true firs and pines (Schmitt and others 2000).
Heterobasidion annosum is a genetically diverse organism
that is composed of at least three intersterility groups (ISGs),
termed S-, P-, and F-types. These groups cannot be distin-
guished based on appearance, but they differ in pathogenicity
and host range and are intersterile (cannot mate) when paired
with each other (Goheen and Otrosina 1998; Sullivan and
others 2001). Only the S-type (which affects mainly true firs,
hemlock, spruce, and Douglas-fir) and the P-type (which
affects mostly pines, larch, incense cedar, and juniper) are
found in the Inland West (Otrosina and Ferrell 1995; Goheen
and Otrosina 1998). Disease-induced mortality of ponderosa
pine, in the Inland West, is often observed on dry sites of poor
quality (Goheen and Goheen 1989; Lockman 1993). Mortality
can also be quite high in natural grand and white fir stands,
especially stands that have experienced selective harvesting
(Schmitt and others 1984; Chase 1989; Goheen and Goheen
1989). Annosus root disease is often associated with mortality
of ponderosa pine or Jeffrey pine in second-generation or
partially harvested stands (Schmitt and others 2000). Specific-
ity of the P- and S-types only applies to living hosts; both types
will colonize dead wood of almost all coniferous species
(Otrosina and Scharpf 1989; Goheen and Otrosina 1998).
Because of differences within and among H. annosum ISGs
and their interaction with host tree species and environmental
factors, the behavior of annosus root disease is highly variable
(Goheen and Otrosina 1998; Garbelotto and others 1999).
Recent research information about annosus root disease has
been compiled in Woodward and others (1998).
The fungus spreads primarily in two ways: aerial spread, and
local underground tree-to-tree spread (Schmitt and others
2000). Basidiospores released from fruiting bodies of the
fungus can travel up to 161 km (100 miles) (Goheen and
Otrosina 1998). Heterobasidion annosum readily colonizes
fresh stumps and stem wounds of all coniferous species except
red and yellow cedar (Wallis and Reynolds 1970; Filip and
Morrison 1998), and spread from colonized stumps can cause
subsequent infections of residual trees via root contacts after
thinning operations (Morrison and Johnson 1978; Sullivan and
others 2001). Stump surfaces are vulnerable to colonization for
up to 4 weeks after harvesting or thinning (Otrosina and Ferrell
1995). However, stump vulnerability to H. annosum basid-
iospores significantly decreases after a period of 2 weeks, as
stumps become colonized by competing fungi. Hyphae from
germinating basidiospores colonize the wood of the stump and
lateral roots. Root grafts or contacts, where roots from two or
more trees touch, can provide contact between infected and
healthy trees and are a source of asexual spread of H. annosum.
Infection centers can often be traced to larger stumps that were
colonized soon after harvest. Stumps greater than 36 cm (14
inches) in diameter are more likely than smaller stumps to
serve as foci of infection centers (Goheen and Otrosina 1998).
Incidence and severity of annosus root disease varies consid-
erably, depending on site location and history of forest stands.
In general, annosus root disease represents a more serious
problem in managed forests than in natural forests (Shaw and
others 1994). Previous studies have shown that higher levels of
mortality are typically found in stands that have been partially
cut and stands that have experienced multiple thinnings (Schmitt
and others 1984). In one survey in Oregon, 89 percent of true
fir stumps were colonized 5 to 9 years after harvest (Filip and
others 2000). In the Inland West, this disease can cause
substantial tree mortality, especially when associated with
drought stress or insect attack (Filip and Morrison 1998). Trees
infected with annosus root disease are frequently subject to
attack by insects, such as the fir engraver beetle on true firs
(Cobb and others 1974; Hertert and others 1975; Lane and
Goheen 1979; James and Goheen 1981; Schowalter and Filip
1993) and mountain pine beetle or western pine beetle on pines
(Cobb and others 1974). In northern Idaho and northeastern
Washington, stands that are dominated by Douglas-fir can
have large amounts of mortality, especially on dry sites
(Lockman 1993; Schmitt and others 2000). Heterobasidion
annosum can persist on a site and within stumps for long
periods. In larger stumps, H. annosum can survive up to 60
years (Goheen and Otrosina 1998).
Disease Symptoms and Signs
Common symptoms of annosus root disease are the same as
for many other root diseases and include yellowing or thinning
of crowns, reduction in tree height and lateral branches, and
stress cone crops. In firs, heartwood in butts and lower stems
of infected trees are often stained red (fig. 7). Advanced decay
includes stringy, spongy, and white-streaked areas that often
have scattered black flecks (fig. 8). Decayed wood can also
become laminated (separated along spring wood of growth
rings) as it dries out.
Fruiting bodies occur in hollow stumps, on root crotches, or
on root collars of dead and dying trees. Although rare or
inconspicuous, fruiting bodies can be used for positive identi-
fication of the fungus. Conks are perennial and may appear to
have several layers or furrows. In the early stages of develop-
ment, these conks appear as small buff-colored cushions. In
later stages, the conks are shelf-shaped, the upper surface is
smooth and a darker brown than the cream-colored margin,
and the undersides are cream-colored with tiny-pores (fig. 9).
A layer of white fungal mycelium (ectotrophic mycelium) is
occasionally seen growing on the exterior of infected roots. In
the absence of other identification methods, wood samples
held at high humidity produce distinctive asexual fruiting
bodies (conidiophores and conidia). DNA-based methods are
12 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Figure 7—Common staining and decay caused by
Heterobasidion
annosum
as it appears in a cross section of hemlock. (Photo
courtesy of W.G.Thies)
Figure 8—White pocket rot characteristic of wood decay caused by
Heterobasidion annosum.
White mycelium with black specks is a common sign
of this disease.
(Photo courtesy of J. W. Schwandt)
13
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Figure 9—Fruiting bodies of
Heterobasidion annosum
,
the cause of annosus root disease, are perennial and
often found in recessed hollows of stumps and roots.
These were observed on an uprooted grand fir. (Photo
courtesy of R.C. Rippy)
also available to help identify H. annosum to species (Garbelotto
and others 1999; Bahnweg and others 2002).
Disease Management
Protective measures have focused primarily on preventing
the colonization of stumps by germinating basidiospores
(Morrison and others 1989; Schmitt and others 2000; Johansson
and others 2002). Chemical and biological treatments of stumps
have been tested with varying levels of success (Driver 1963a,b;
Edmonds and others 1969; Weir 1969; Driver and others 1970;
Russell and others 1973a; Nelson and Li 1980; Woodward and
others 1998). One boron-containing compound (sodium
tetraborate decahydrate) has been widely used and can be
successful for preventing colonization if applied within 48
hours after harvesting (Otrosina and Ferrell 1995; Schmitt and
others 2000). Alternatively, in Europe, freshly cut stumps may
be treated with urea, as this increases the pH of the stump
surface to a level where basidiospores cannot survive (Brantberg
and others 1996; Johansson and others 2002).
The potential for biological control of H. annosum has been
the subject of limited investigations in Western North America
(Filip and Morrison 1998). A fungal species, Phlebiopsis
gigantea (=Peniophora gigantea), has been documented to
out-compete H. annosum when applied to freshly-cut stumps
(Rishbeth 1963; Woodward and others 1998). In Europe, use
of this biological agent for controlling H. annosum is wide-
spread, commercially available, and effective (Vainio and
others 1998).
Stump removal can be effective for reducing buildup of root-
rot fungi after tree removal; however, this method is expensive
and can be disruptive to the site (Thies and others 1994; Roth
and others 2000). Because H. annosum can behave as a wound
parasite, wound prevention, especially during harvest, is a
14 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
primary method of disease management (Goheen and others
1980; Aho and others 1983; Filip and others 1992b; Sullivan
and others 2001). Heterobasidion annosum populations vary
in their pathogenicity and host ranges, and hosts also vary in
their ability to resist strains of the pathogen (Worrall and others
1983). This pathogen can be extremely damaging on sites
planted with a single species. Thus, one approach used to
manage annosus root rot is favoring mixed stands that include
hardwoods or a mixture of conifers (Schmitt and others 2000).
Timing of management activities may be important, but
results from different studies are in conflict. Thinning in
summer or winter may be preferred over spring or fall. Thin-
ning under warm temperatures may allow other fungi such as
Trichoderma viride to out-compete H. annosum for establish-
ment within the substrate of newly created stumps (Driver and
Ginns 1969). Alternatively, if thinning occurs during winter
when snow cover is present or temperatures are at or below
freezing, the rate of stump colonization by H. annosum may be
reduced and may favor colonization by other microorganisms
(Chernykh and Belyi 1978). Studies in northeastern Oregon,
however, found no difference in stump colonization by H.
annosum among different harvesting seasons (Filip and others
1992a).
Fuels Treatment Considerations
Thinning may be one of the most important management
practices that encourages the spread of H. annosum because it
opens infection routes, creates fresh stumps for colonization by
H. annosum, and creates logging wounds on standing trees that
serve as infection sites for H. annosum (Aho and others 1983;
Korhonen and others 1998; Sullivan and others 2001). On the
other hand, one study showed that thinning in western hemlock
stands increased sound-wood volume in residual trees despite
minor logging wounds and H. annosum-caused decay (Goheen
and others 1980). Also, wound prevention and stump treatments
have been shown to greatly reduce H. annosum infection.
In pine stands in the Southeastern United States, annosus
root disease has been reduced by prescribed burning prior to
thinning operations. This burning reduces the duff layer that
favors sporophore production, thus reducing the spore loads in
the stand at the time of thinning (Froelich and others 1978;
Otrosina and others 1995). However, successful tests of
underburning for control of annosus root disease have not been
reported for forests in the Inland West, perhaps because this
treatment is typically considered impractical under the stand
and soil conditions in this region. Research on the effects of
prescribed fire on the incidence of stump-surface infection is
needed in the Inland West.
15
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Laminated Root Rot _____________
Laminated root rot, caused by Phellinus weirii (Murr.) Gilb.,
is prevalent in British Columbia, Washington, Oregon, north-
ern California, northern Idaho, and western Montana (Nelson
and others 1981; Sturrock and Garbutt 1994; Thies and Sturrock
1995). Phellinus weirii is considered a “disease of the site”
because it persists across tree generations (Hadfield and others
1986; Thies and Sturrock 1995). This disease can cause exten-
sive growth loss and mortality that is often associated with
windthrows within disease centers involving several conifer
species of diverse age classes. The pathogen appears to spread
by contact of susceptible tree roots with stumps or roots of
previously infected trees. Laminated root rot is most destruc-
tive in stands 25 to 125 years old, and damage increases in
subsequent stands (Tkacz and Hansen 1982).
In Western forests, highly susceptible species include Dou-
glas-fir, grand fir, mountain hemlock, and white fir. Interme-
diately susceptible species include Engelmann spruce, Pacific
yew, subalpine fir, western hemlock, and western larch. Spe-
cies that are tolerant or resistant to laminated root rot include
lodgepole pine, sugar pine, western white pine, ponderosa
pine, and western redcedar; all hardwoods are immune (Filip
and Schmitt 1979; Hadfield and others 1986; Nelson and
Sturrock 1993; Thies and Sturrock 1995). Fire exclusion
increases the proportion of shade-tolerant conifers, which are
generally more susceptible to laminated root rot.
Phellinus weirii has two basic forms: one that produces a
butt rot in western redcedar, and another that causes root
disease in Douglas-fir and several other conifer species (Thies
and Sturrock 1995). A proposal to change the scientific name
of the form of P. weirii affecting Douglas-fir to P. sulphurascens
(Larsen and Cobb-Poulle 1990) or Inonotus sulphurascens
(Larsen and others 1994) has not been universally accepted.
Phellinus weirii often occurs with other root diseases, such as
Armillaria root disease and black stain root disease (Sinclair
and others 1987). More than 4.4 million cubic meters (154
million cubic feet) of timber volume are estimated to be lost
annually to laminated root rot (Nelson and others 1981).
Associations have been found between the occurrence of P.
weirii and aspect, elevation, timber type, habitat type, and soil
type. Soil characteristics, such as pH, temperature, and mois-
ture content, may also affect the occurrence and extent of
damage from laminated root rot (Thies and Sturrock 1995).
Mortality due to this disease is generally higher on wetter than
on drier habitat types.
Trees infected with P. weirii are sometimes killed by bark
beetles in combination with other root diseases. The Douglas-
fir beetle, Douglas-fir pole beetle, and fir engraver are com-
monly associated with laminated root rot (Schowalter and Filip
1993). Laminated root rot, as with other root diseases, provides
a continuous source of suitable material for maintaining en-
demic bark beetle populations when conditions are not favor-
able for epidemics (Thies and Sturrock 1995).
Disease Symptoms and Signs
Infected trees exhibit crown symptoms typical of many root
diseases: chlorosis, reduction in growth, gradual crown thin-
ning, and a crop of stress cones. In wind-thrown trees, decayed
roots are typically broken at the root collar; roots do not usually
lift from the soil as the tree falls (fig. 10). This is especially
noticeable in infection centers, where fallen trees may occur as
a random pattern of crossed stems. Incipient decay in the roots
and lower bole of infected trees usually gives a red-brown stain
to the outer heartwood (fig. 11). Advanced decay separates
easily into sheets at the annual rings; sheets contain small pits
and are covered with reddish-brown to pinkish setal hyphae
that can be seen with a hand lens (fig. 12). Gray-white to buff-
colored mycelia may be found on the outer surface of infected
roots. Fruiting bodies of P. weirii typically form near or in
contact with the forest floor on the underside of downed trees
and uprooted stumps. They have light gray-brown pore sur-
faces and margins that are white to cream-colored when young
but later turn a uniform chocolate brown. However, fruiting
bodies are of little diagnostic value, as they are uncommon and
inconspicuous (Hadfield and others 1986; Sinclair and others
1987).
Disease Management
Identifying areas containing laminated root rot is the essen-
tial first step to successfully managing for this disease. After
disease has been assessed, acceptable levels of disease must be
determined before deciding upon a management response. To
reduce disease impacts, thinning treatments can:
•Avoid areas with the disease.
•Selectively remove the most susceptible species.
•Favor root-disease resistant or tolerant species.
•Make openings in which to plant or regenerate root
disease-resistant or tolerant species.
•Use push-felling to extract root systems from the soil
(Bloomberg and Reynolds 1988; Morrison and others
1988).
•Include stump removal to reduce inoculum levels
(Hadfield 1985; Hadfield and others 1986; Thies and
Sturrock 1995).
•Include a combination of these treatments.
However, the potential adverse effects of soil compaction
and tree wounding caused by heavy equipment must also be
considered (Lull 1959; Froehlich and McNabb 1984; Smith
and Wass 1991). Thus far, effects of treatments such as
fertilization have supported increased growth of trees but
produced no apparent reduction in disease-caused tree mortal-
ity (Thies and Nelson 1988). Eradication of P. weirii from
infected stumps with fumigants (Thies and Nelson 1987) in
conjunction with antagonistic Trichoderma (Nelson and Thies
1985, 1986; Nelson and others 1987) has been successful
under experimental conditions but has not been used opera-
tionally in the United States. Increasing species diversity to
16 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Figure 10—Laminated root rot mortality center showing characteristic root
wads with roots broken near the root collar, leaving only stubs. (Photo
courtesy of W.G. Thies)
Figure 11—Incipient decay from laminated root rot in the roots and lower bole of infected trees is frequently
observed as a red-brown stain in the outer heartwood. (Photos courtesy of W.G. Thies)
17
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
reduce losses associated with P. weirii is a preferred strategy
in northern Idaho, western Montana, Oregon, and Washington
(Hadfield and others 1986; Thies and Sturrock 1995; Filip
1999).
For additional information pertaining to P. weirii and its
management, see Morrison and others (1992), Thies and
Sturrock (1995), and Hansen and Goheen (2000).
Fuels Treatment Considerations
The effect of wildfire on P. weirii has been speculated to be
more indirect than direct, as fire acts by changing stand species
composition and forest successional patterns involving suscep-
tible and resistant hosts (McCauley and Cook 1980). Because
fire has little direct influence on the survival of P. weirii
Figure 12—White mycelium on the outer bark of infected root underneath the duff is a common characteristic of laminated root rot
(upper left). Advanced decay from laminated root rot causes wood to separate easily along growth rings of spring-wood. Each layer
of decayed wood has small pits and often displays cinnamon-brown mycelium with hairlike pegs that project away from the layer
(upper right). The underside of this Douglas-fir log shows the appressed habit of the young fruiting bodies of
Phellinus weirii
(lower). (Photos courtesy of W.G. Thies and R.C. Rippy).
inoculum, fuels reduction by burn treatments, such as pre-
scribed burning, are not likely to directly influence the
occurrence of laminated root rot (Thies and Sturrock 1995).
However, fuels treatments that would use prescribed burning
at intervals that approximate historic fire frequencies, with the
aim of converting stand composition back to an early succes-
sional stage, are strategies for management of this disease.
From a production standpoint, thinning is generally not recom-
mended in stands with a major component of susceptible trees
and moderate to high levels of disease. In disease centers,
thinning treatments without stump removal may increase dis-
ease severity. This approach is generally only considered when
either a final harvest is scheduled within 15 years, or severe
losses are deemed acceptable (Thies and Sturrock 1995).
18 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Black Stain Root Disease _________
Black stain root disease is caused by three host-specialized,
genetically isolated varieties of the pathogen Leptographium
wageneri (Kendrick) Wingfield (Harrington and Cobb 1984,
1986, 1987; Zambino and Harrington 1989, 1990). The hard
pine variety (var. ponderosum) is prevalent from the Pacific
Coast States to the Northern Rockies, including all States and
Provinces of the Inland West south and west of the Bitterroot
Mountain range. This variety infects ponderosa, lodgepole,
and Jeffrey pine and can also occasionally be found in hem-
locks, white and Engelmann spruce, and white pines (Morrison
and Hunt 1988; Allen and others 1996). The Douglas-fir
variety (var. pseudotsugae) is broadly adapted to Douglas-fir
from moist coastal to arid situations, from California to British
Columbia to southern Colorado and New Mexico. The pinyon
variety (var. wageneri) occurs on pinyon species, primarily
singleleaf pinyon, throughout the Southwest to the southern
edge of Idaho.
Stands affected by black stain root disease are generally
moist to mesic. Disease incidence may be higher in areas with
soil compaction, poor drainage (Hadfield and others 1986), or
coarse-textured soils and shallow soils prone to summer drought
(Morrison and Hunt 1988). The pinyon and hard pine varieties
are limited to areas with cool soil temperatures, whereas the
Douglas-fir variety can survive and infect at higher soil and
bole temperatures. Black stain root disease occurs as persistent
infection centers or as scattered infections after disturbances
such as road building, construction, and thinning (Hansen
1978; Harrington and others 1983). Within infection centers,
the pathogen spreads by growth in and along roots, by root
grafting or root contact, and by limited hyphal growth (up to 15
cm) through the soil between roots (Cobb 1988). Infection can
occur through natural openings in small roots, such as splits at
root forks and sites where lateral roots emerge, or at the site of
root injuries, such as caused by abrasion against rocks (Hessburg
and Hansen 2000). Root lesions from Armillaria infections on
adjacent trees can also allow transfer of the black stain root
disease pathogen from host to host (Hessburg and Hansen
2000). Root damage caused by equipment can provide an
abundance of infection courts. Experiments with inoculated
Douglas-fir seedlings indicate that most minor root wounds no
longer act as infection courts after 1 month, because wound
closure prevents infection. The pathogen does not penetrate
either intact parenchyma or callus (Hessburg and Hansen
2000).
Expansion of infection centers in the various hosts averages
around 1 m/year (3.3 feet/year) (Hessburg and others 1995),
with rates of up to 7 m/year (23 feet/year) reported in some
pinyon stands (Cobb 1988). Even the slower rate is substan-
tially (three to five times) faster than most other root diseases
(Hessburg and others 1995). It appears that the black stain
pathogen may be active in stumps for only 3 years after harvest,
as it dies rapidly after the death of host tissues (Cobb 1988).
Severity and rates of spread are positively related to density of
hosts (Cobb 1988). In Douglas-fir, rates of infection center
expansion decrease markedly after stands reach an age of 30 to
35 years (Hessburg and others 1995).
Unlike many root pathogens, infection centers of black stain
root disease are established from initial infections vectored by
root-feeding bark beetles and weevils (Goheen and Cobb
1980; Harrington and others 1985; Witcosky and Hansen;
1985; Witcosky and others 1986; Cobb 1988; Hansen and
others 1988; Harrington 1988). Insect vectors are attracted to
stressed trees (Harrington and others 1985; Morrison and Hunt
1988). Trees in decline due to black stain root disease are also
predisposed to infestation by both root-inhabiting and bole-
inhabiting species of bark beetles (Goheen and Cobb 1980;
Cobb 1988; Hessburg and others 1995, 2001; Negron and
Wilson 2003). Bark beetle species that vector the disease are
attracted to volatiles given off by living roots of dying trees
(Witcosky and others 1987; Kelsey and Joseph 1998). Once an
insect has caused a tree to become infected, recent layers of
xylem in the root system are colonized. Presence of persistent
black stain root disease infection centers can maintain local
populations of bark beetles, initiating bark beetle mortality
centers (Cobb 1988).
Disease Symptoms and Signs
Crown symptoms that occur if black stain root disease is
present in a stand include many typical above ground symp-
toms of root disease, such as reduced growth and fading of
crowns, slow or rapid decline (slower in old trees and faster in
young trees), distress cones, and crown wilt. Pines will usually
succumb to bark beetle attacks before the latter symptom can
be expressed (Cobb 1988). Because mortality can be hastened
by bark beetles, and the crown symptoms of black stain root
disease are similar to those of other root diseases, mortality
caused by black stain root disease is sometimes mistakenly
attributed to other causes (Cobb 1988; Hessburg and others
1995). However, stands with black stain root disease will have
trees in different stages of mortality, typical of root disease
infection centers, instead of a uniform onset of mortality,
typical of mass attacks by bark beetles (Cobb 1988).
The most diagnostic sign of black stain root disease is the
microscopic mycelium that is visible as dark purple to black
staining in younger layers of sapwood in roots and the lower
trunk of an affected tree (fig. 13). Unlike blue stain fungi,
which are also vectored by insects, these streaks of intense
staining in the bole strictly follow the path of xylem upward
from the roots where the initial infection point occurred. In
cross-sections of stems and roots, the staining has an arclike
appearance, as it lies primarily within early wood of recently
infected growth rings. This staining pattern further differenti-
ates black stain root disease from blue stain, which develops in
parenchyma rays at the time of mortality and extends in wedge-
shaped patterns into heartwood (fig. 14). The restriction of
black stain to recent rings results from fungal colonization of
the tracheids of active xylem (Cobb 1988). These symptoms
and the development of disease are similar to the vascular wilts
of hardwoods (Joseph and others 1998).
19
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Figure 13—Dark purple to black streaks of stain within the outer one to two annual rings of
sapwood in the lower bole and roots of an affected tree indicate black stain root disease caused
by
Leptographium wageneri
. (Photos courtesy of W.G. Thies and R.C. Rippy)
Figure 14—Arcs of stain (left) are typical of black stain root disease and are caused by
Leptographium
wageneri
colonizing the tracheids of the most active growth rings. In contrast (right), fungi that cause blue
stain colonize parenchyma rays to form wedgelike sections that penetrate to the limits of the sapwood.
(Photos courtesy of W.G. Thies and R.C. Rippy)
20 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Disease Management
Few long-term field studies have been conducted to assess
the behavior of this disease over time in stands under different
types of management. Nonetheless, indications from existing
studies suggest that the disease may be minimized by maintain-
ing stand vigor while avoiding disturbance and injury, espe-
cially during times of peak beetle activity (Hadfield and others
1986; Hessburg and others 1995). Maintaining trees in vigor-
ous condition with adequate spacing has been suggested as the
most useful long-term strategy for minimizing losses to black
stain root disease (Cobb 1988).
Because black stain root disease is vectored by beetles, any
activities that increase the numbers of vectors in an area have
the potential to increase disease, if disease is already present in
stands within flight distances of its vectors. The distance for
concern is 1.6 km (1 mile) for insect vectors of Douglas-fir
(Hansen and others 1988; Hessburg and others 1995). Black
stain root disease has long been associated with road building
(Goheen and Hansen 1978; Hansen 1978; Hessburg and others
2001) and thinning-associated disturbance (Harrington and
others 1983; Hessburg and others 2001). Thinning of affected
species can trigger increases in vector population and estab-
lishment of many new infection centers. Dramatic 20- to 100-
fold increases in the most important of the three insect species
that act as vectors in Douglas-fir have occurred following
springtime precommercial thinning (Harrington and others
1985; Hansen and others 1988). Thinning and clearcutting in
ponderosa pine in winter and spring also cause dramatic
increases in vector beetles (Otrosina and Ferrell 1995). Man-
agement strategies for Douglas-fir have included preventing
tree wounds, avoiding tractor logging within 1.6 km (1 mile) of
an infection center, favoring a diverse stand composition that
includes tree species that are not susceptible to the pathogen,
and restricting thinning to summer after spring beetle flights
(Hansen and others 1988; Filip 1999; Hessburg and others
1995, 2001). This ensures that vectors will be established in
other dead or dying trees prior to thinning. Summer thinning
also allows roots and stumps of newly cut trees time to dry out
before fall (Witcosky and others 1986; Hessburg and others
1995; Filip 1999). Fall thinning is not recommended because
stumps may still attract insects the following spring. A specific
recommendation for minimizing disease by the hard pine
variety of the pathogen is to maintain mixed conifer type
forests where indicated by habitat, but limit susceptible pine
species to 25 percent of the mix when disease is present (Cobb
1988). If stands of susceptible tree hosts have adequate spacing
and are intermixed with nonsusceptible species, these stands
will be at lower risk of becoming permanent infection centers
and will have reduced tree-to-tree spread through roots. Be-
cause the pathogen has low persistence, healthy but susceptible
trees outside the margins of infection centers have been felled
to limit further expansion (Tainter and Baker 1996).
Fuels Treatment Considerations
Factors to consider prior to choosing a fuels treatment are
whether black stain root disease occurs within the vicinity of
the treatment area, which hosts are being affected, the ratio and
spacing of host and nonhost trees, and the choice of season for
fuels management treatments. In general, if a fuels treatment is
necessary, the key for minimizing subsequent black stain root
disease is to reduce wounding of residual trees, minimize
compaction, and time activities in relation to insect dispersal
activities. Little published information is available on the
effects of prescribed fire in different seasons on host vigor and
root diseases (Thies and others 2001). Research is needed to
determine the degree to which insect vectors of black stain root
disease are attracted to roots damaged by prescribed fire,
whether the damage or attraction differs with the season during
which the fire treatment is applied, and how long insect
attraction persists. Thus, answers are likely to be complex, and
must weigh the direct detrimental effects on hosts against the
potential effects on disease and fire behavior. A study of
ponderosa pine/juniper stands in the Blue Mountains of south-
eastern Oregon showed that all root- and lower bole-feeding
insect species that could potentially act as vectors of black stain
root disease were greater in abundance following prescribed
burns than in unburned controls. Fall burning caused greater
direct and delayed mortality than spring burning, but mortality
was not due to black stain root disease (Thies and others 2001;
Thies and others in press). Different species of root- and lower
bole-feeding insects were favored by spring or fall burns, and
no clear relationships between time-of-burn or transmission of
black stain root disease had been identified at these sites (Thies
and others 2001). Another study in the same location showed
that fall burning caused significant reductions in both fine root
biomass and ectomycorrhizal fungus species richness in stands
of ponderosa pine, both important for tree health, whereas
spring underburning had little effect (Smith and others 2004).
21
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Schweinitzii Root and
Butt Rot _______________________
Phaeolus schweinitzii (Fr.) Pat., also known as the velvet top
fungus, causes Schweinitzii root and butt rot, a decay in the
roots and lower stem that affects most mature coniferous tree
species in Western forests (Hadfield and others 1986). It is the
most common cause of brown-cubical rot in Douglas-fir and
other commercial timber species (Gilbertson and Ryvarden
1987). Douglas-fir is the most common species infected, but
ponderosa pine, lodgepole pine, western white pine, western
redcedar, western larch, western hemlock, spruces, and true
firs are also known hosts (Allen and others 1996; Hagle and
others 2003). Phaeolus schweinitzii is regarded as the most
serious butt-rot pathogen of conifers in old-growth forests in
the United States (Sinclair and others 1987).
Windborne spores of P. schweinitzii colonize trees through
basal scars caused by fire, wounds from logging, or through
wounded roots. The pathogen might also spread between trees
with root contact, but little direct evidence is available to
support this theory (Sinclair and others 1987). This fungus
usually infects and kills root tips, resulting in “stubbed” roots
with blunt ends (Hagle and others 2003). Soil compaction,
drought, and flooding may predispose roots to damage (Tainter
and Baker 1996). Trees growing on sites with shallow soils or
soils with low water-holding capacity are more likely to be
damaged by P. schweinitzii (Beckman and others 1998).
Phaeolus schweinitzii can grow through wood colonized by
Armillaria species and may predispose trees to Armillaria root
disease and bark beetles (Hadfield and others 1986; Sinclair
and others 1987). Alternatively, Armillaria species may be a
secondary pathogen that follows P. schweinitzii in Douglas-fir
and grand fir in Idaho (Dubreuil 1981).
Decay is usually confined to the roots and lower 3 m (9.8
feet) of the bole. Trees with advanced Schweinitzii rot are
predisposed to windthrow and breakage. Valuable butt logs are
extensively decayed, causing serious losses in timber volume.
Decay caused by P. schweinitzii may be confused with decay
caused by Fomitopsis pinicola, Fomes officinalis, Laetiporus
conifericola, and Postia sericeomollis (Partridge and others
1978; Allen and others 1996; Hagle and others 2003).
Disease Symptoms and Signs
Trees infected with P. schweinitzii rarely exhibit root-dis-
ease crown symptoms (Partridge and others 1978; Hadfield
and others 1986; Hagle and others 2003). Carpenter ant infes-
tation and/or bark beetle attacks may occur on trees weakened
by this disease. Small-diameter roots that are infected may be
“stubbed” with gall-like swellings and may have dark, resinous
heartwood (Hagle and others 2003). Infected trees may also
have pronounced swelling at the butt (Scharpf 1993).
Fruiting bodies are annual, developing in the late summer
and fall, but may persist for a year or more (fig. 15). These
fruiting bodies are found on the ground near infected trees or
occasionally emerge directly from the stem of an infected tree
(Hadfield and others 1986; Hagle and others 2003). Those that
form on the ground are circular or irregularly lobed in form
with a sunken center and short, central stalk; those forming on
infected trees are shelflike. The upper surface is velvety, up to
25 cm (10 inches) in diameter, and usually reddish brown with
a yellowish margin when fresh. The lower surface has large
angular pores and is yellow-green when young, turning brown
with age. Older fruiting bodies can resemble cow dung in
appearance (Scharpf 1993; Hagle and others 2003). Incipient
decay, though sometimes inconspicuous, often produces a
yellow to red-brown discoloration in the heartwood. With
advanced decay, wood becomes reddish brown, crumbly, and
may crack into large cubical pieces. Decayed wood can be
easily crumbled into fine powder. Thin, white mycelial mats
are sometimes present in shrinkage cracks (Gilbertson and
Ryvarden 1987; Hagle and others 2003).
Disease Management
To reduce infection by P. schweinitzii, care should be taken
to avoid wounding of trees and minimize soil compaction
during harvesting operations. Because Schweinitzii rot prima-
rily impacts older trees, economic losses might be reduced by
harvesting trees before they are overmature (Hadfield and
others 1986). Because of possible windthrow (fig. 16), trees
infected with P. schweinitzii in recreation areas and around
buildings should be considered hazardous. Increment boring
of the lower bole and roots of trees can be used to detect brown
rot. At other sites, the degree of care used to prevent new
disease may be less. However, the beneficial ecological roles
of brown-rots should also be considered before selecting
management activities. For example, brown-rotted logs once
incorporated into the forest floor or mineral soil can act as
moisture reserves for ectomycorrhizal root development in
periods of drought, provide a source of N fixation, and/or
improve soil physical properties (Harvey and others 1987;
Page-Dumroese and others 1994; Jurgensen and others 1997).
Fuels Treatment Considerations
Prescribed fire hot enough to cause basal wounds on trees
should be avoided. Infections can be reduced by minimizing
tree wounding during thinning operations. Information is
unavailable for other potential interactions of fuels treatments
with Schweinitzii root and butt rot.
22 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Figure 16—Trees infected with
Phaeolus
schweinitzii
may break off above ground
level, exhibiting characteristic brown cubical butt rot (left).
Phaeolus
schweinitzii
decay 30.5 cm (12 inches) above ground level in fire scar (right). (Photos courtesy
of R.C. Rippy and J.D. Rogers).
Figure 15—Fruiting bodies of
Phaeolus schweinitzii
form on the
ground or cut stumps of affected trees. They are circular or
irregularly lobed in form, with a sunken center and short, central
stalk. Older fruiting bodies, such as in the lower photo, dry up and
closely resemble cow dung—hence the common name “cow-pie
fungus.” (Photos courtesy of J.E. Stewart and W.G. Thies)
23
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Other Root Diseases of
Consideration in the
Inland West ____________________
Other notable root diseases present in the Inland West are
Tomentosus root disease, Rhizina root rot, and stringy butt rot.
While Tomentosus root disease, also known as stand-opening
disease, has been the subject of numerous studies in British
Columbia and Ontario, less information has been reported on
it for the Inland West. Published information is also quite
limited for Rhizina root disease and stringy butt rot.
Tomentosus Root Disease
Tomentosus root disease is found throughout the temperate
region of the northern hemisphere (Hunt and Unger 1994). In
Oregon and Washington, Engelmann spruce is the most com-
mon host (Hadfield and others 1986). Lodgepole pine is an
important host in other parts of the range, while true firs,
western white pine, and Douglas-fir are occasional hosts
(Hagle and others 2003). The fungal pathogen, Inonotus
tomentosus (Fr.) Teng., primarily spreads slowly from host-to-
host by root-to-root contact but it also spreads by spores
(Lewis 1997). Because of the slow rate of spread, infection
centers occur as clumps of dead trees in the landscape, with
standing dead or windthrown trees surrounded by trees in
various stages of disease progression (Lewis 1997).
The likelihood that Tomentosus root disease will occur on a
site depends on species composition, stem density, and site
characteristics, such as soil moisture and nutrient regime
(Lewis 1997; Bernier and Lewis 1999). Tomentosus root
disease is difficult to control once I. tomentosus has become
established at a site. The fungus can utilize root and stump
resources well after bole removal, allowing it to persist at least
30 years after timber harvest. In a study in Utah, I. tomentosus
was isolated from 75 percent of stumps 20 years after cutting.
It was also isolated from dead roots as small as 1.3 cm (0.5
inches) in diameter 9 years after cutting and up to 5.5 m (18
feet) away from an infected stump 20 years after cutting (Tkacz
and Baker 1991). Postharvest burning did not reduce the
incidence of I. tomentosus, as the fungus was recovered in the
same proportion of stumps from burned and unburned areas.
Disease Symptoms and Signs—Classic root rot symptoms,
including thinning crown, chlorotic foliage, and reduced height
growth, are displayed by trees infected with I. tomentosus
(Bernier and Lewis 1999). Crown symptoms may not develop
until 50 percent of a tree’s roots are infected, so it is difficult to
recognize Tomentosus root disease until after it is well estab-
lished (Filip 1986; Lewis 1997). Characteristics of the decay in
the root and butt heartwood are diagnostic. Appearance ranges
from a reddish-brown stain in the early stages of decay, to
elongate, spindle-shaped white pockets separated by firm
reddish-brown wood in advanced decay (Tkacz and Baker
1991; Hagle and others 2003). In spruce, disease caused by I.
tomentosus can be differentiated from annosus root rot, be-
cause wood decayed by the former is firm, instead of spongy.
Also, the annual conks of I. tomentosus, which are produced on
the ground near infected trees, are readily distinguished from
the perennial conks of H. annosum. Fruiting bodies of I.
tomentosus are 5 to 12.7 cm (2 to 5 inches) in diameter, with
velvety caps and short stems. The pore surface on the underside
of the cap is yellow and cream-colored, and darkens with age
(Hagle and others 2003).
Rhizina Root Rot
Rhizina root rot is a disease of burned sites that affects
conifer seedlings. This disease is not responsible for large
volume losses, but it can impact plantings and natural regen-
eration after fire (Ginns 1968; Morgan and Driver 1972).
Spores of Rhizina undulata Fr.:Fr. are viable for up to 2 years,
and spore germination is stimulated by fire (Gremmen 1971).
After underlying or adjacent soils have been warmed by fire,
pathogen spores will germinate quickly and colonize the
recently sterilized soil. The pathogen may infect susceptible
conifer seedlings and cause mortality within a growing season.
Infection occurs when a susceptible conifer root contacts
mycelium or strandlike rhizoids. Losses of up to 80 percent of
planted stock have occurred (Callan 1993); however, an exten-
sive survey of 277 burned clearcuts in western Washington and
Oregon found that Rhizina root rot caused little damage (Thies
and others 1979). Rhizina root rot does not occur at all sites
exposed to fire. For this disease to occur, sites must have the
pathogen (usually from past infections), acidic soils, and
susceptible conifer roots. In a laboratory study, young seed-
lings of lodgepole pine, a fire-successional conifer species,
displayed strong disease symptoms when exposed to an isolate
of Rhizina undulata from British Columbia (Egger and Paden
1986). While the pathogenicity of this fungus was consistent
and intense, field occurrences either go unrecognized or occur
infrequently.
Disease Symptoms and Signs—Seedling symptoms are
nonspecific and may be easily misidentified as other root
diseases or drought. The presence of Rhizina fruiting bodies in
the vicinity of a dead conifer seedling is reliable evidence of the
pathogen’s presence on the site. Fruiting bodies of this fungus
are brown to black, crustlike and fleshy, slightly convex, and
often have an undulating margin. The undersides of the fruiting
bodies have root-like rhizoids at multiple locations, which
attach them to the soil, host roots, and occasionally to other
substrates in the soil. They can occur in groups or lines that
follow buried substrates or parasitized roots (Ginns 1974).
Stringy Butt Rot
Perenniporia subacida (Peck) Donk. produces a white stringy
butt rot in infected trees and may also be known as yellow root
rot because of its distinctively colored mycelium (Hadfield and
others 1986). It is most commonly associated with mature trees
that are either suppressed or weakened (Hadfield and others
1986). True firs, western hemlock, western redcedar, Engel-
mann spruce, and white spruce are among the most susceptible
of its many coniferous and deciduous hosts. Perenniporia
24 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
subacida has been classified as a wound parasite, although it is
unclear whether infection by spores or mycelium is primarily
responsible for disease initiation (Zeglen 1997). Trees infected
by P. subacida are subject to windthrow and mortality. Signifi-
cant volume loss from stringy butt rot is not an important issue
in the Inland West (Hadfield and others 1986).
Disease Symptoms and Signs—Initially, the decay appears
as a light-brown heartwood stain. Distinctive yellow mycelial
mats may be observed when sheets separate along the annual
rings; these mycelial mats help identify the fungus as P.
subacida. Progression of the decay leads to small white pits in
the sheets of annual wood, but eventually only white spongy
fibers with black flecks remain. Leathery, perennial fruiting
bodies that are cream to yellow in color may occupy the
underside of downed wood or the basal region of standing dead
trees. Small, circular pores mark their surface (Gilbertson and
Ryvarden 1987).
Disease Management and Fuels Treatment
Considerations
Tomentosus Root Disease—The ability of I. tomentosus to
colonize organic debris and remain on site for decades is an
important characteristic of this root disease. The survival and
spread of I. tomentosus in an infected stand may be exacerbated
by stumps and dead roots created by thinning practices. These
reservoirs of inoculum may never contact susceptible trees if
susceptible spruce and pine species are not planted. By delay-
ing the regrowth of susceptible species for 20 years after tree
removal, roots and stumps may decay to a water-soaked
condition that reduces the likelihood of new infections. At least
one study indicates that I. tomentosus cannot be recovered
from such material (Tkacz and Baker 1991). In some situa-
tions, stump removal may be a viable option. Thinning meth-
ods that minimize wounding of residual trees would also limit
the establishment of new infections. Prescribed burning to
reduce woody substrate would not likely kill all of the fungus
that resides within root heartwood.
Standing or windthrown trees dead from or having
Tomentosus root disease contribute to fuel loading and may
provide substrate for spruce beetles. One study reported that
diseased trees were not predisposed to attack and may have
helped to maintain spruce beetle populations at low levels
(Lewis and Lindgren 2002). Infected trees can live 15 to 20
years before mortality occurs. The gradual decay of the roots
makes infected trees subject to windthrow and a hazard con-
cern for recreation areas.
Rhizina Root Rot—Even without seedlings to infect and
parasitize, Rhizina undulata has the ability to use woody debris
as a nutrient source. The fungus Rhizina may actually aid the
decomposition of some smaller organic debris (Callan 1993).
Described as only weakly saprotrophic, it is not known how
long the fungus can remain on site without a living host.
In the past 15 years, no new research has been published on
Rhizina root rot in the Inland West, so little is known about the
interaction of fuels treatments, fire, and Rhizina root rot. If
susceptible conifer seedlings were to be planted, it would be
important to reduce spore germination by minimizing the areas
of soil subject to warming. One possible strategy would be the
use of “pile and burn” practices for removing hazardous fuels.
More studies are needed to determine the scope of this disease
in the Inland West and appropriate fuels treatments on sites
with this disease.
Stringy Butt Rot—With so many hosts susceptible to
infection by P. subacida, the best methods to manage stands
infected with stringy butt rot are to prevent the creation of
wounds that facilitate new infections on healthy trees and to
remove diseased trees during thinning. The effects of fuels
treatments and fire on stringy butt rot have not been well
studied. Not enough is known about the distribution of this
disease and the extent of its impact on the Inland West.
Additional research studies are needed to determine impacts of
fuels treatments on this disease.
25
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
Summary of Fuels Management
Considerations _________________
Before prescribing fuels treatments, managers should
know whether root diseases are present and where they are
located on a site. For conifer forests of the Inland West, root
diseases caused by A. ostoyae, H. annosum, Phellinus weirii,
L. wageneri, and Phaeolus schweinitzii are of special concern
because of the extreme damage and mortality they can cause.
The interactions between trees, root pathogens, and environ-
ment are complex and vary by forest structure, stand history,
habitat type, species composition, soil characteristics, bark
beetle populations, and activity of other forest insects and
pathogens. It is not uncommon to have more than one root
disease present in a stand.
Fuels treatment options should consider past management
practices and their effects on current stand conditions. Seral
species tend to be tolerant of root diseases but have declined in
many areas of the Inland West. In northern Idaho, the com-
bined effects of blister rust, selective harvesting, planting
practices, and fire exclusion have reduced stand representation
of white pine, western larch, and ponderosa pine but increased
those of root disease-susceptible Douglas-fir and true firs
(Beckman and others 1998; Filip 2002). On sites with mixed
species composition—where seral and climax species co-
occur with root disease organisms—infection and mortality of
susceptible species have the potential to prolong the seral
phase. This can favor the seral species that are tolerant to root
disease unless late seral or climax species have been favored by
management (Meyer 2004).
Use of a single type of treatment will not be appropriate for
all forest conditions and root diseases. Some fuels treatments
may increase root disease incidence, so managers should
decide at the outset of planning what levels of mortality and
growth loss will be acceptable. Estimates of treatment effects
on mortality will be important for making subsequent esti-
mates of changes in surface and ladder fuels. Tree mortality
may be attributed directly to the root disease or to indirect
consequences of infection, such as susceptibility to bark beetle
attack. Killed trees can persist as standing snags or as
windthrows, and may increase the overall amount of woody
debris. Thus, fuels reduction treatments that cause increases in
root disease mortality will likely increase the accumulation of
fuels.
Acceptable options for fuels treatments differed among the
diseases included in this paper. Thinning may increase damage
from some root diseases, such as Armillaria root rot, laminated
root rot, and annosus root disease. Incidence of black stain root
disease and Schweinitzii root and butt rot may be less affected
by this management practice if specific guidelines are fol-
lowed. For all root diseases, care should be taken to avoid
wounding of trees during timber harvesting or prescribed fire
to reduce future impacts on stand health. Thinning treatments
should also favor the species most tolerant to root disease and
be timed to avoid problems with bark beetles. For example,
regarding annosus root disease, biological and chemical con-
trol applied at the time of thinning could be effective for
preventing new infections. Little information is available on
the effects of prescribed fire and when it should be applied
(seasonally) to decrease the effect of most root diseases.
Despite the importance of these issues, research regarding
interactions of fuels treatments with root diseases has been
limited. For this reason, much of the information presented
here has been based on general principles of forest pathology.
More precise information for specific sites, diseases, and host
trees would be desirable to optimize fuels treatments with
stand conditions and locations. To determine the most suitable
forest management practices for the Inland West, more studies
are needed on the interactions among fuels treatments, fire, and
root diseases.
26 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-141. 2005
References _____________________
Aho, P.E.; Fiddler, G.; Filip, G.M. 1983. How to reduce
injuries to residual trees during stand management activities.
Gen. Tech. Rep. PNW-156. Portland, OR: U.S. Department
of Agriculture, Forest Service, Pacific Northwest Forest and
Range Experiment Station. 17 p.
Allen, E.A.; Morrison, D.J.; Wallis, G.W. 1996. Common tree
diseases of British Columbia. Victoria, BC: Natural Re-
sources Canada, Canadian Forest Service, Pacific Forestry
Centre. 178 p. Available online at http://www.pfc.forestry.ca/
diseases/CTD/.
Bahnweg, G.; Möller, E.M.; Anegg, S.; Langebartels, C.;
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