Bark Beetle Outbreaks in Western North America: Causes and Consequences

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Beetle Bark
Symposium,
Snowbird Utah,
November 2005
Bark Beetle Outbreaks in
Western North America:
Causes and Consequences

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i
Bark Beetle Symposium Participants
Snowbird, UT, November 15 – 18, 2005
Barbara Bentz
,
Editor and Co-organizer
, USFS
Rocky Mountain Research Station
Jesse Logan
,
Co-organizer
, USFS Rocky Mountain Research Station
Jim MacMahon
,
Facilitator
, Ecology Center, Utah State University
Craig D. Allen
, USGS, New Mexico
Matt Ayres
, Dartmouth College
Ed Berg
, USFWS, Alaska
Allan Carroll
, Canadian Forest Service, British Columbia
Matt Hansen
, USFS Rocky Mountain Research Station
Jeff Hicke
, University of Idaho
Linda Joyce
, USFS Rocky Mountain Research Station
Wallace Macfarlane
, GeoGraphics, Inc., Utah
Steve Munson
, USFS Forest Health Protection
Jose Negrón
, USFS Rocky Mountain Research Station
Tim Paine
, University of California at Riverside
Jim Powell
, Utah State University
Ken Raffa
, University of Wisconsin
Jacques Régnière
, Canadian Forest Service, Quebec
Mary Reid
, University of Calgary
Bill Romme
, Colorado State University
Steven J. Seybold
, USFS Pacific Southwest Research Station
Diana Six
, University of Montana
Diana Tomback
, University of Colorado, Denver
Jim Vandygriff
, USFS Rocky Mountain Research Station
Tom Veblen
, University of Colorado
Mike White
, Utah State University
Jeff Witcosky
, USFS Forest Health Protection
David Wood
, University of California at Berkeley
Staff
Hannah Nordhaus
, writer
Mitchelle Stephenson
, design

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
ii
Acknowledgements
We thank Stephanie White, Paige Weed, Ryan Davis, Donovan Gross, and Greta
Schen for assistance at the workshop and in manuscript preparation. Neil Cobb,
Dave Breshears, and Cynthia Melcher provided helpful comments on the manu-
script. For assistance with aerial overflight data of bark beetle impacts we thank
Jeanine Paschke, Gurp Thandi, and GeoGraphics, Inc. Elizabeth Grossman
assisted with an early version of the manuscript. Funding was provided by the
USDA Forest Service Rapid Science Assessment Team.

S
ince 1990, native bark beetles have killed billions of trees across millions of
acres of forest from Alaska to northern Mexico. Although bark beetle infesta-
tions are a regular force of natural change in forested ecosystems, several of the
current outbreaks, which are occurring simultaneously across western North
America, are the largest and most severe in recorded history.
There are many species of bark beetles, but only a few are responsible for
the large areas of dead trees we see today. These species colonize live and
recently downed trees, and kill either the entire tree or a portion of it dur-
ing colonization and brood production. Bark beetle ecology is complex and
dynamic, and a variety of circumstances must coincide for a bark beetle
outbreak to succeed on a large scale. Only when specific conditions are
met—ranging from suitable climatic conditions across an entire region, to
the particular forest structure and age, to the existence of certain bacteria
and fungi within the beetle and host tree—will bark beetle populations
grow large enough to infest and kill trees across large landscapes.
Although outbreak dynamics vary from species to species and from forest to for-
est, the combination of two major factors appears to be driving the current out-
breaks.
• Changing climatic conditions, in particular elevated temperatures and
drought: elevated temperatures can speed up bark beetle reproductive
and growth cycles and reduce cold-induced mortality during cold snaps.
Extreme or prolonged water stress, caused by a combination of drought
and warm temperatures, can weaken trees, making them more suscepti-
ble to bark beetle attacks.
• Forest history and host susceptibility: many conifer forests in western
North America contain dense concentrations of large, mature trees that
are highly susceptible to bark beetle outbreaks. The factors that have
contributed to these conditions vary in relative importance from area to
area. They include large stand-replacing fires (both natural and human-
set) and timber harvesting near the end of the 19th century—distur-
bances that resulted in large areas of forest of similar size and age. In
some areas, fire suppression over the last century also has inhibited the
growth of new trees and created more dense stands. Because aggressive
bark beetles favor mature trees, older, even-aged stands that have
regrown or been replanted after disturbance events such as wildfire and
harvesting may be more vulnerable to future bark beetle outbreaks
than younger, more diverse stands. When trees must compete for
resources in crowded conditions caused by either natural or human
processes, bark beetles can more easily overcome stressed trees’ defens-
es and initiate a severe outbreak.
Report Highlights
1
A bark beetle outbreak
requires:
• Abundant suitable host
trees
• A climate favoring bark
beetle survival and
development

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
Given the complexity of bark beetle community dynamics and the specific ecosys-
tems they inhabit, the roles these factors play differ from forest to forest.
Although research has uncovered a great deal of information about the life cycles
and host interactions of some species of bark beetles, many gaps in our knowl-
edge remain. In addition, because changing climate and forest disturbances have
altered outbreak dynamics in recent years, some of what has been learned from
past outbreaks may no longer hold true. There may be no equivalent in the 100
or so years of recorded history for the current outbreaks.
These recent infestations may result in dramatic changes to the long-term eco-
logical pathways of some ecosystems, radically shifting vegetation patterns in
some hard-hit forests. The visual landscape cherished by many nearby landown-
ers and visitors is altered as well, as once-green trees turn brown and then lose
their needles. In addition, bark beetle-killed trees pose some hazards when dead
trees fall in areas of forest that humans frequent.
Although there are no known management options to prevent the spread of a
large-scale bark beetle outbreak, land-use activities that enhance forest hetero-
geneity at the regional scale—such as creating patches of forest that contain
diverse species and ages of trees—can reduce susceptibility to bark beetle out-
breaks. However, because resource objectives often differ, and because the factors
influencing a bark beetle outbreak vary depending on the species, host tree, local
ecosystem, and geographical region, there is no single management action that is
appropriate across all affected forests.
2
A whitebark pine forest in Yellowstone National Park. The red trees were attacked and killed
by mountain pine beetles the year before the photo was taken in July 2007.
PHOTO BY JANE PARGITER, ECOFLIGHT, ASPEN CO

3
IN THIS REPORT:
Bark Beetle Ecology and Biology .............................................................................................6
Aggressive Bark Beetle Species, Hosts, and Habitats..............................................6
Life Cycle and Life Stages of a Bark Beetle ...............................................................8
Bark Beetles and Temperature ..................................................................................10
Bark Beetle Flight and Pheromone Plumes ............................................................12
Bark Beetles and Trees ................................................................................................14
The Community Beneath the Bark ..........................................................................15
Bark Beetle Outbreaks..............................................................................................................17
Historical Bark Beetle Outbreaks ..............................................................................18
Are the Current Outbreaks Different? ......................................................................20
Contributing Factors.................................................................................................................23
Climate.............................................................................................................................23
Disturbance and Human Influence............................................................................26
The Future of Our Forest Ecosystems .................................................................................28
Ecological Consequences of Recent Bark Beetle Outbreaks ...............................28
Bark Beetles and Fire ...................................................................................................33
Research Gaps................................................................................................................38
Additional Reading ...................................................................................................................38

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
Are the current bark beetle outbreaks unprecedented?
Because it is technically difficult to reconstruct bark beetle outbreaks before the late 1800s, we are
uncertain how extensive or severe outbreaks may have been prior to that time. However,
scientists have examined outbreak frequency and severity over the last few centuries by
examining tree-ring growth patterns, and made inferences about bark beetle presence in forest
ecosystems prior to the last few centuries using ancient pollen and bark beetle remains. They also
use computer models that describe how bark beetles respond to temperature to analyze beetle
behavior in the past and compare it to current outbreaks. In doing so, they hope to answer two
questions.
1) Is the scale of the current outbreaks—in terms of both geographic extent and the diversity
and number of affected ecosystems—different from previously recorded outbreaks?
2) Have the underlying dynamics and mechanisms of bark beetle systems been altered because
of new inputs to the forest systems in which bark beetles live? For instance, have a warming
climate, air pollution, and/or historical patterns of forest management and fires altered forest
habitat and beetle dynamics, allowing beetle populations to explode in recent years?
4
T
ravel through a western North American forest today, and you will probably
notice large areas of standing dead trees with dry reddish-brown needles, or
ghostly gray snags from which all the needles have fallen. Across the West, from
Alaska to Canada, throughout the Rocky Mountain region and the southwestern
United States, many forests and hillsides are now blanketed with trees that have
been recently infested and killed by various species of bark beetles.
These outbreaks of aggressive bark beetles, which are occurring in numer-
ous forest ecosystems across western North America, are the biggest in
recorded history. The term “aggressive” describes those species that can kill
either the entire tree or a portion of it during colonization and brood pro-
duction. Although western forests have experienced regular infestations
throughout their history, the current outbreaks are remarkable for their
intensity, their extensive range, and their simultaneous occurrence in mul-
tiple ecosystems across the continent.
These beetles are not only attacking forests where they have traditionally
been found, but they also are thriving in some places where widespread
infestations have not previously been recorded. Some outbreaks reflect the
expansion of at least one bark beetle species beyond its recorded historical
range.
With so many forests severely affected, land owners, land managers, policymak-
ers, and the general public have taken notice. The extraordinary extent of the
outbreaks has prompted concern that this massive loss of trees may impair
ecosystem functioning and reduce the ability of our forests to provide future
wildlife habitat, to protect watershed quality, to store carbon, and to be a source
of timber and recreational opportunities.
Introduction
New outbreaks of
mountain pine
beetle in British
Columbia and
Alberta are a sign
of expansion
beyond the
beetle’s recorded
historical range.

Why are these unusual bark beetle
outbreaks occurring across western
North America, and what will they do
to our forests? In November 2005,
scientists with the USDA Forest
Service, Rocky Mountain Research
Station convened a conference in
Snowbird, Utah. Entomologists, ecol-
ogists, and foresters from across the
continent shared the latest research
on bark beetle outbreaks and sought
to explain the causes, historical con-
text, and short- and long-term conse-
quences of the current outbreaks.
This publication is a product of that
workshop. It first explains how bark
beetles function within their native
ecosystems, and then examines the
recent outbreaks and explains how they differ from previous recorded infesta-
tions. Finally, the report explores the ecological effects of the current outbreaks
and identifies areas where more research may be needed so we better under-
stand the causes and consequences of current and future infestations.
5
The current bark beetle outbreaks differ from previously recorded
infestations because of:
Their intensity — bark beetles are killing trees in larger numbers, at a
faster pace, and over longer time periods
Their extent — bark beetle outbreaks are occurring in numerous
ecosystems from Alaska to northern Mexico
Their synchroneity — bark beetle outbreaks are occurring concurrently
across western North America
Mountain pine beetle-killed lodgepole pine.

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES Bark Beetle Ecology and Biology
N
ative insects, including bark beetles, are among the greatest forces of natural
change in forested ecosystems of North America. Every few decades, depend-
ing on weather and local forest conditions, bark beetle populations increase and
infest large areas of conifer forest. In doing so, they play an essential role in
forests’ natural cycle of growth and
regeneration.
Bark beetles don’t just kill trees; they
also have beneficial effects. For example,
certain lodgepole pine cones require heat
from forest fires, which are sometimes
fueled by beetle-killed trees, to release
the seeds within. In addition, because
beetles generally attack larger trees,
they serve an essential regenerative
function—they help renew forests by
killing older and declining trees, allow-
ing younger, more productive trees to
compete successfully for light, nutrients,
and water.
Aggressive Bark Beetle Species, Hosts, and Habitats
Of the hundreds of different North American bark beetle species, only a handful
of species (fewer than one percent) is considered aggressive, meaning the beetles
kill all or a portion of the host trees they infest. These few species are primarily
responsible for the large areas of tree mortality we see across the major forest
ecosystems of western North America, from the spruce forests of the far North to
the pine forests of the southwestern U.S.
6
These photos show the same Tenaya Creek drainage in Yosemite National Park. The photo on
the left was taken in 1925, five years after a 1920 mountain pine beetle outbreak. The photo
on the right, from 1984, shows how forests regenerate after beetle attacks.
PHOTO A BY J.M. MILLER PHOTO B BY M. FURNISS
A Source of Forest Renewal
Historically, bark beetles have not “destroyed” forests; rather,
they have served as positive forces of transformation that
redistribute nutrients and growing space. Whereas older trees
often die en masse during bark beetle outbreaks, younger
trees are usually not attacked. Released from competition for
light, nutrients, and water, the young trees grow quickly to
replenish the forest canopy. Thus, bark beetles can help
mature forests regenerate. However, our understanding of
the role bark beetles play in forests is based on observations
from the past few centuries. As climate and local ecosystems
change, the balance between bark beetles and their host
forests also may change.

The tree-killing bark beetles reside in a single family of insects (Curculionidae,
subfamily Scolytinae), and each species has evolved to feed and reproduce in a
single conifer group (Table 1). The Douglas-fir beetle, for instance, is found exclu-
sively in Douglas-fir trees across western North America. The mountain pine
beetle attacks and reproduces in at least 12 different species of North American
pine across a number of ecosystems, from sea level to 10,000 feet in elevation,
from the Pacific coast to the Black Hills of South Dakota, and from Baja
California to central British Columbia. Its range has historically been limited by
climate, not by the availability of host trees. For instance, although lodgepole
pine trees are found as far north as northern British Columbia, western Alberta,
and the Yukon and Northwest Territories, those areas until very recently had
been untouched by mountain pine beetle. In recent years, however, mountain
pine beetle outbreaks have been recorded in northern British Columbia and
western Alberta; lodgepole pine forests in the Yukon and Northwest Territories
still appear to be free of bark beetles.
7
TABLE 1.
A g g ressive Bark Beetles and their Host Trees in the We s t e rn U.S. and Canada
mountain Dendroctonus lodgepole pine, ponderosa pine, bristlecone
pine beetle ponderosae pine, whitebark pine, western white pine,
sugar pine, limber pine, and others
spruce beetle Dendroctonus Engelmann spruce, white spruce,
rufipennis Lutz spruce, Sitka spruce
Douglas-fir beetle Dendroctonus Douglas-fir
pseudotsugae
western pine Dendroctonus ponderosa pine,
beetle brevicomis Coulter pine
southern pine Dendroctonus Apache pine,
beetle frontalis Chihuahua pine,
ponderosa pine
Arizona fivespined Ips lecontei ponderosa pine
ips
piñon ips Ips confusus piñon pine
western balsam Dryocoetes confusus subalpine fir
bark beetle
fir engraver Scolytus ventralis white fir, California red fir, grand fir
COMMON NAME SCIENTIFIC NAME MAJOR HOST TREE SPECIES IN THE U.S AND CANADA
PHOTO BY DANIEL RYERSON
Adult piñon ips beetles are only 0.12 to 0.14
inches long—barely larger than a grain of
rice. Mountain pine beetles are slightly larg-
er, around 0.20 inches, while spruce beetles
average 0.22 inches long.

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
Species within the genus
I p s
, such as the piñon ips and
Arizona fivespined ips, also can kill their hosts, although typi-
cally they are not considered major disturbance agents. In
recent years, however, elevated population levels of a number
of
I p s
species have coincided with drought, resulting in large
areas of mortality, particularly in piñon and ponderosa pine
forests of the southwestern U.S.
Life Cycle and Life Stages of a Bark Beetle
Native North American bark beetles in the genera
Dendroctonus, Dryocoetes, Ips,
and
Scolytus
live, feed on,
and lay their eggs in the soft living tissue of a tree, known as
the phloem, which lies just below the tree’s protective outer
bark layer. These small beetles are 1/8th to 1/4 inch long,
dark brown to black, and cylindrical in shape. Bark beetle
larvae, which develop from eggs laid inside the tree, are
white and less than 1/4 inch long, thus resembling a grain of
rice. After maturing through four life stages—egg, larva,
pupa, and adult—they emerge as new adults from the host
tree and fly to a new live tree, where they mate and lay eggs
to begin the cycle again (Figure 1).
Bark beetle life cycles range from a few weeks to two or three
years, depending on the species and climate in which the
beetles are found. Adult beetles attack their host trees during
8
Ponderosa pines generally are found at lower elevations.
Whitebark pines are found
at higher elevations in the
northern Rocky Mountains
of the U.S. and in the
Canadian Rocky Mountains,
as well as in Oregon,
Washington, and California.

the time of year when they are most vul-
nerable—for example, spruce, true fir,
and Douglas-fir trees are less able to
fend off a beetle attack when their roots
are still under snow. Bark beetles typical-
ly attack these trees in late spring, when
air temperatures have warmed but snow
remains on the ground.
Once a beetle finds and successfully bur-
rows into a new tree, it builds chambers
or shallow tunnels,
known as “galleries,” in
which the female bee-
tles lay their eggs. Each
female can lay up to
200 tiny white or cream-colored
eggs, which hatch within one to
three weeks. After egg hatch,
the larvae tunnel around the cir-
cumference of the tree, feeding
as they go and impeding the
flow of water and nutrients
between the tree’s roots and nee-
dles in the crown. Larvae of
some species (e.g.,
southern pine beetle
and western pine bee-
tle) tunnel into the
outer bark to complete
development. Fungi
introduced into a tree
by bark beetles also
may aid in tree death.
Although the needles
of an infested tree remain green for months after
the initial attack, once a sufficient number of
attacking adult beetles have initiated galleries
and laid eggs, the tree has little chance of surviv-
ing. Following attack, the tree’s needles turn yel-
low, orange, then red, and finally brown over a
one- to two-year period. Eventually the needles
drop to the forest floor. The length of time the
dead tree remains standing depends on tree
9
FIGURE 1.
Life Stages of a Bark Beetle
Bark beetle eggs.
Most adult bark
beetles chew a
vertical or winding
egg gallery in an
infested tree and
then lay their
eggs in individual
niches along the
length of the
gallery.
Mountain pine beetle larvae burrow horizontally
across the host tree, eating phloem as they go.
Other bark beetle species have winding galleries.
Bark beetle pupae. The pupal
resting state is an intermedi-
ate stage that each beetle
passes through as it goes
from larva to adult.
Bark beetles develop through four stages—egg, larva, pupa, and
adult, before emerging to fly and attack another tree.

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
species and locale.
A typical beetle-
killed lodgepole
pine will fall to
the ground within
five to 10 years.
Decomposition
takes place more
slowly in extreme-
ly arid climates—
a number of high-
elevation whitebark pines killed during a late 1920s mountain pine beetle out-
break in central Idaho, for instance, still remain standing today.
Bark Beetles and Temperature
Bark beetles, like all insects, are “ectotherms,” and thus their development, sur-
vival, and reproductive success are highly temperature-dependent. Temperature
influences everything in a bark beetle’s life, from the number of eggs laid by a
female to the beetle’s ability to disperse to new host trees.
All western bark beetles have specific, evolved responses to temperature that can
differ dramatically from species to species, and even within species. Most
research on temperature-dependency in western North American bark beetles
has been conducted with mountain pine beetle and spruce beetle. Although the
general relationships described here may apply to other species, the specific
details apply only to these two species.
10
Whitebark pine killed by mountain pine beetle, most likely in the late
1920s.
Trees turn reddish-orange
within a year after bark
beetle infestation, then
brown and eventually gray
as the needles fall to the
ground.

From early in a bark beetle’s
life, temperature regulates the
rate of development, determin-
ing when the insect moves
from one life stage to the next.
Different species—and even
different populations within
species with large geographical
ranges—respond to different
temperature thresholds.
Mountain pine beetles, for
instance, move into the next
life stage only when enough
thermal energy has accumulat-
ed and temperatures surpass a
certain set threshold, which
varies depending on location.
Those in the northern portion
of the mountain pine beetle’s
range develop faster, and move
into the pupal and adult life
stages only when temperatures
have risen above 15 degrees
Celcius (C). This reduces the
risk of cold-induced mortality
and ensures emergence during
summer months when air tem-
peratures have warmed, and
when some species of host trees are more vulnerable to attack. In southern cli-
mates, pupation appears to occur at cooler temperatures, an adaptation that syn-
chronizes populations with the warmer environment. Other species, such as the
spruce beetle, maintain synchrony by going into a resting state called “diapause,”
and resume their development only after detecting specific cues, such as temper-
ature thresholds and/or longer periods of daily warming of the tree bark by solar
radiation.
11
FIGURE 2.
Mountain Pine Beetle
Temperature Thresholds for
Development in Central Idaho
Mountain pine beetles develop to the next life stage
only when enough thermal energy has accumulated
and temperatures reach a certain threshold. These
temperature thresholds and thermal energy require-
ments differ depending on geographic location, and
help the beetles avoid cold-induced mortality during
the sensitive larval and pupal stages. They also help to
synchronize adult emergence.
Temperature and Beetles
Temperature influences everything in a bark beetle’s life, from how it develops, to how quickly it
reproduces, to how long it lives. Colder temperatures keep beetle populations in check through cold-
induced mortality and longer reproductive cycles. When temperatures warm, beetle populations in
colder climates thrive.
Temperature in degrees (C)

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
Temperature is also a limiting factor in moun-
tain pine beetle survival. Low temperature
extremes, especially at certain times of the
year when beetles are particularly sensitive,
can result in widespread beetle mortality. To
reduce cold-induced losses, the beetles’ devel-
opment is timed to progress through the egg
and pupal stages, the most cold-susceptible
stages, during warmer weather (Figure 3).
The insects also generate “antifreeze” com-
pounds that circulate through their bodies and
help them survive in colder locales. Although
bark beetles are well-adapted to cold winters,
unusual cold snaps in the spring and fall may
occur when antifreeze concentrations are low,
thus resulting in widespread mortality.
Finally, temperature influences how quickly
beetles complete a generation, and thus how
often they reproduce. Whereas many bark bee-
tle species that live in warmer, lower-elevation forests can develop numerous
generations in a single year (“mulitvoltine”) or one generation in a single year
(“univoltine”), beetles that live at higher elevations, where weather is harsher
and warm periods are shorter, most often reproduce on a less frequent “semivol-
tine” schedule, producing one generation every two years. Recent increases in
temperature and growing-season length at higher elevations, however, have
shifted many semivoltine, cold-weather-adapted beetles to a univoltine life cycle,
resulting in rapid population growth.
Warming temperatures, combined with exceptionally susceptible host trees of
similar age classes or weakened by crowding, drought, or old age, are believed to
be a significant driver of large spruce beetle outbreaks in Alaska, Utah, and
Colorado. Warmer temperatures also appear to be a factor in mountain pine bee-
tle outbreaks in lodgepole pine forests of British Columbia and Colorado, and in
high-elevation white pine forests throughout the northern and central Rocky
Mountains.
Bark Beetle Flight and Pheromone Plumes
As described above, temperature determines the developmental timing of each life
stage beneath the bark. Temperature also plays a role in the timing of adult
emergence from trees and beetle flight. Once summer temperatures have warmed
to approximately 16 °C to 18 °C, newly developed adults of most species begin a
brief period of maturation feeding and then chew through the bark and fly to a
new tree, synchronizing their emergence to attack their new hosts en masse.
12
FIGURE 3.
Phloem Temperature of Lodgepole Pine
in Central Idaho
Mountain pine beetle live and feed in pine-tree phloem, a thin layer
that lies just beneath the outer bark. The beetles live within the phloem
for an entire year, and survive exposure to sub-freezing temperature s
in the fall, winter, and spring by generating “antifreeze” compounds.
Aug 25 Oct 14 Dec 3 Jan 22 Mar 13 May 2 June 22
30
20
10
0
-10
-20
-30
-40
——— Maximum daily phloem temperature
——— Minimum daily phloem temperature

Beetles find their host trees based on a combination of factors: some appear to land
randomly on new trees, whereas others seem to select their hosts based on either a
visual assessment of size, or certain odors. Because larger trees provide an easier
target and often have thicker phloem that allows female beetles to produce more
offspring, aggressive bark beetles are more likely to attack large trees than small,
young trees.
I p s
species are an exception, often attacking and killing small trees.
Although the exact mechanism is not known, scientists suspect that beetles most
likely fly randomly until they encounter a “pheromone plume” that has been
released by “pioneer” beetles that have selected a specific tree for attack. The
process of chewing through the bark and ingesting tree phloem causes the syn-
thesis of an attractive odor (pheromone) within the insects’ guts. This odor can
attract thousands of beetles to the same tree within a few days. As more beetles
attack, more pheromone is produced. This is critical to the success of bark beetle
attacks, as large numbers of beetles are required to overwhelm a tree’s defenses.
Once the tree is saturated with adult beetles, the attractive pheromone dissi-
pates, repellant odors may be produced, and the tree no longer attracts addition-
al beetles.
There are a number of factors that determine how far the beetles travel. Bark
beetles prefer dense stands with tree canopies that touch, keeping attractant
pheromones under the canopy where beetles can detect them. By flying below
the tree tops but above the underbrush, they can best encounter pheromones
emitted by other beetles. If there are no suitable host trees in a stand, the bee-
tles will disperse farther and move into new areas to find susceptible trees. If the
stand is very open and receives more heat and sunlight, the pheromones dissi-
pate. Without pheromone attractants within the stand of trees, the beetles move
up above the canopy, where they are often picked up by wind drafts and carried
passively into other areas.
Wind currents can carry beetles above the forest canopy for many miles. For
instance, recent infestations in northern Alberta, where 20th century mountain
pine beetle outbreaks have not been recorded previously, are believed to have
been started by beetles carried on upper air currents from infest-
ed areas in British Columbia. In the U.S., transport of infested
logs from bark beetle-infested areas to those without active popu-
lations also may contribute to the long-distance movement of
bark beetles.
Bark Beetles and Trees
How does a tiny beetle kill something as big as a tree?
Through sheer numbers: the more beetles that attack a tree, the
easier it is to kill it. Trees do have defenses against bark beetle
attacks. They produce a sticky, sap-like substance called resin
13
Overcoming a tree’s defenses
Trees have natural defenses against
beetle attacks. These can be overcome
when:
1. trees are weakened by drought,
disease or excessive competition
2. a large beetle population develops
in the vicinity and attacks en
masse, overwhelming even
healthy trees

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
that contains toxic defensive chemicals. If a tree is healthy and vigorous and the
number of beetles attacking is small, the tree can produce enough resin to kill the
attacking beetles and fend off the attack. These defensive mechanisms, however,
can be exhausted if many beetles attack the same tree within a few days.
Although many trees may be “suitable” for bark beetle infestations, not all are
“susceptible” to attack. The most suitable trees—those whose thick phloem pro-
vides the best food for developing larvae—are generally healthy trees that have
received adequate water, light, and nutrients. They often can resist the
onslaught of a relatively small number of bark beetles. Susceptible trees, by con-
trast, are often unhealthy and easily overcome by even a small
number of beetles, but often do not provide a good resource for
larval feeding and survival. A number of factors—drought, tree
diseases, and overcrowding—can make trees more vulnerable to
beetle attacks, and in some cases bark beetles may infest weak-
ened trees already doomed to die.
In general, dense, older stands, where trees must compete for
resources, are more susceptible to bark beetle outbreaks, where-
as heterogeneous landscapes that contain many sizes, ages, and
species of trees are more resistant and resilient. Because less
crowded forests foster healthier trees, and because increased
sunlight and wind in open tree canopies help disperse the
pheromones that encourage mass bark beetle attacks, some stud-
ies indicate that removal of trees, or thinning, may reduce sus-
ceptibility in some forest types.
Even the healthiest tree, however, can not withstand a mass
attack, and the explosion in beetle numbers in recent years has
made extensive outbreaks more likely, killing healthy and
unhealthy trees alike. Although beetles rarely kill all of the trees
in a forest, it may appear that way in the months before the
14
In 1997, an unusual
wind event on the
Routt National Forest
blew down large
swaths of Engelmann
spruce across 25,000
acres. Spruce beetle
populations multiplied
in the downed trees
and then attacked
green trees outside of
the blowdown area.
Trees respond to bark beetle attack by releasing a resin that is toxic to the invading beetles. If the tree
is vigorous and few beetles are attacking simultaneously, it can survive an attack.

beetle-killed trees’ discolored needles drop. In several recent outbreaks, however,
more than 90 percent of suitable host trees were killed by bark beetles in some
stands.
Large numbers of recently downed trees also can contribute to severe outbreaks of
some bark beetle species, particularly the Douglas-fir beetle and spruce beetle.
When live trees are blown down, their phloem can remain suitable for bark beetle
development up to a year later, whereas the trees’ resinous defenses are depleted
quickly due to the loss of contact between roots and soil. Beetle population num-
bers increase within the downed trees, and emerging adults can then successfully
attack surrounding live host trees. An unusual wind event in 1997 in spruce
forests on the Routt Divide in Colorado created
the largest known blowdown in the Rocky
Mountain region, and spruce beetle population
levels increased dramatically in some of the
affected areas.
The Community Beneath the Bark
Upon colonizing a tree, each beetle brings
with it an array of fungi, nematodes, bacteria,
and mites. Although the ecological roles of
many of these organisms are not well under-
stood, it is known that most bark beetles are
symbiotically associated with specific fungi
that may be important for beetle survival.
Mountain pine beetles, for example, transport
two fungal species in specialized body structures called mycangia (Figure
4). These fungi travel with the beetles when they disperse, and then
grow in the phloem and sapwood of trees attacked by the beetles. The
fungi may aid in successful colonization of a tree, and developing beetle
larvae and adults also feed on the fungi, which help the beetles to
process tree nutrients and also may provide vital nutrients not found in
tree phloem. Research suggests that the fungi are also critical for brood
development. Mountain pine beetles developing with fungi are larger than those
without fungi, a size difference that can mean an increase in egg production of
up to 300 percent.
Natural enemies and competitors of bark beetles also are attracted to bark bee-
tle-infested trees. A wide variety of predators and parasitoids exploit bark bee-
tles’ pheromone communication system to find and feed on beetles walking on
the outer bark and within trees they have infested. These natural enemies,
which include checkered beetles and many species of woodpeckers, help keep
normal, low-level bark beetle populations in check. Other less aggressive phloem-
and sapwood-feeding beetle species also may compete with aggressive bark bee-
tle species for food resources.
15
Adult moun-
tain pine bee-
tles feed on
fungal spores
prior to
emergence
from the
brood tree.
Many bark beetle species
introduce symbiotically associ-
ated fungi into trees as they
enter through the bark. The
growing bark beetle larvae
feed on the fungi, which give
the phloem and sapwood a
bluish tint.

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
Over time, trees killed by
bark beetles also are invaded
by a number of other organ-
isms, including wood-boring
insects and rotting fungi,
which are important in the
tree’s decay process. Once
bark beetles have infested a
tree, that tree becomes a ver-
itable jungle of hitchhiking, predatory, and decay
organisms.
Many bark beetle associates, in particular insect
natural enemies and fungal species, are also sensi-
tive to temperature, developing and reproducing
when temperatures reach specific thresholds. It is
not clear how climate warming may change the relationship between beetles and
their hitchhiking fungal, predatory, and parasitic associates.
16
Three-toed woodpeckers feed on bark
beetles and also make nests in trees
the beetles kill.
PHOTO BY RYAN BRACEWELL
PHOTO BY VICKI SAAB
A checkered beetle eat-
ing an adult mountain
pine beetle.
FIGURE 4.
Bark Beetle Fungal Associates
Scanning electron micrographs showing the opening of the mycangia (My) of a mountain
pine beetle, which is located on the maxillary cardine (C) near the mouth. The scanning
electron micrographs show fungal material (F) protruding from the mycangia.
PHOTOS BY KATHY BLEIKER AND SARAH POTTER
CC
F
F
My
My
F
My
My

Bark Beetle Outbreaks
W
ritten and tree-ring records dating back as far as 1750 indicate a consis-
tent record of intermittent outbreaks of aggressive bark beetles in various
forest types across the West. Over the past 10-15 years, however, the frequency,
severity, and extent of bark beetle outbreaks have increased. Bark beetles have
killed large numbers of trees in multiple forest ecosystems throughout their his-
torical range, and also in some ecosystems where outbreaks have never or sel-
dom been recorded (Figure 5).
Bark beetles are active in a number of dif-
ferent species of trees.
• Mountain pine beetles are killing
large numbers of lodgepole pine in
the Rocky Mountain region.
• Mountain pine beetles have been found
in high elevation white pines, such as
whitebark, bristlecone pine, and limber
pine. These pines are generally long-
lived, and are restricted to higher ele-
vations throughout the West. They are
considered foundation species, which
play extremely important roles in the
ecosystems where they are found and
are vital to the survival of other local
s p e c i e s.
• Mountain pine beetle outbreaks are
occurring in northern lodgepole pine
and lodgepole/jack pine hybrid
forests in Alberta.
• Unusually large outbreaks of spruce
beetles are occurring in Alaska,
Utah, Washington, Wyoming,
Arizona, Colorado, and the Yukon
Territory.
• In the southwestern United States,
Ips spp
. beetles have killed extraordi-
nary numbers of ponderosa and
piñon pines.
• Western pine beetles have killed
large numbers of ponderosa pine in
southern California and throughout
the southwestern U.S.
17
FIGURE 5.
Bark Beetle Outbreaks
in Western
North America
Native bark beetles have affected more than 150 million acres of western
North American forests during the past 10 years.
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
Year
35
30
25
20
15
10
5
0
SOURCE: NATURAL RESOURCES CANADA, CANADIAN FOREST SERVICE;
USDA FOREST SERVICE, FOREST HEALTH PROTECTION

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
• Douglas-fir beetle, fir engraver, and western balsam bark beetle are active
throughout the West.
• Southern pine beetle is active in pine forests of southern and central
Arizona.
Since 1997, bark beetles have collectively killed
billions of trees across millions of acres of forest
in western North America. The fact that so
many regionwide bark beetle events are hap-
pening concurrently at such intensity across so
many ecosystems is truly remarkable and sug-
gests common factors.
Historical Bark Beetle Outbreaks
Are the current bark beetle attacks unprece-
dented? To understand the significance of the
current outbreaks, scientists are using a vari-
ety of techniques to piece together the frequen-
cy, intensity, timing, area, and duration of pre-
vious beetle outbreaks and other forest distur-
bance events. Understanding bark beetles' his-
torical range and variations in behavior over
long periods of time is key.
We know that bark beetles have been associat-
ed with forests since at least the Holocene era,
which began approximately 12,000 years ago,
and probably much longer. Remains of
Dendroctonus
from 8000 years ago, for
instance, were identified in lake sediment cores
taken in the Bitterroot Mountains of Montana.
More recently, forest rangers, land managers,
and forest entomologists have recorded out-
breaks of many species throughout the West.
These early written records date back to the
late 1880s and are qualitative in nature. For
example, a forest ranger riding his horse across
vast acres of forest might have noted that bee-
tles had killed a large number of lodgepole pine
in a certain drainage or forest.
Acquiring quantitative assessments of previ-
ous bark beetle outbreaks is more complicat-
ed. It is far more difficult to trace the history
18
Over the past 120 years, forest rangers, land
managers, and forest entomologists recorded
outbreaks of many bark beetle species throughout
the West. In 1934, Chief Ranger George F. Baggley
and Forester Maynard Barrows wrote about a
severe outbreak in Yellowstone National Park:
“The mountain pine beetle epidemic is
threatening all of the white bark and
lodgepole pine stands in Yellowstone Park.
Practically every stand of white bark is
heavily infested...and will be swept clean in a
few years. If the insects spread from the
white bark pine to the lodgepole stands, it
seems inevitable that much of the park will
be denuded.”
- from “Montana’s Thirty Year Mountain
Pine Beetle Infestation,” James C. Evenden, 1944.
Yellowstone Chief Ranger
George T. Baggley survey-
ing a bark beetle-killed
forest near Hellroaring,
Yellowstone National
Park, 1930 (Furniss and
Renkin 2003)
A lodgepole pine “ghost
forest” killed by mountain
pine beetles near Lake
Tenaya, Yosemite National
Park, 1916. (Furniss and
Wickman 1998)
PHOTO BY J. E. PATTERSON

of a bark beetle outbreak than it is to understand fire history in many forest
types. Although forest fires often spare many trees in a forest, leaving behind
scars on surviving trees that scientists can detect in tree rings many years
l a t e r, trees that have been mass-attacked by bark beetles do not live to tell the
tale. It can be difficult to examine the cores of many species of beetle-killed
trees, particularly in areas where decomposition occurs rapidly. In some dry,
high elevation sites where decomposition occurs slowly, however—such as in
high-elevation whitebark pine forests—scientists have been able to use tree
rings of well-preserved, beetle-killed trees to date mountain pine beetle out-
breaks as far back as the late 1880s.
In most instances, scientists must date outbreak events by looking for more sub-
tle evidence in the rings of those trees that survived. Because bark beetles kill
larger trees first, smaller trees in the vicinity of a beetle outbreak and/or non-
host tree species often grow at a dramatically increased rate when freed from
competition for light and water from larger trees nearby. Thus, a sudden and
sustained increase—or “growth release”—in tree-ring width among a number of
trees in a stand during the same period is an indication that the trees survived a
major disturbance event. After factoring out the influence of climate—such as
increased precipitation, which also can result in a wide growth ring—a bark bee-
tle outbreak can be deduced.
Studies using these techniques suggest that spruce beetle and mountain pine
beetle outbreaks have been regular occurrences in western North American
forests in recent centuries. Tree ring analyses on the Kenai Peninsula in Alaska,
FIGURE 6.
Growth Release in Trees Surviving Bark Beetle Outbreaks
19
Trees that survive bark beetle outbreaks often experience a “growth release” — a period of much
faster growth that results in a doubling of tree-ring width over a ten-year period — after sur-
rounding trees have been killed by bark beetles. Historical outbreaks can be dated by measuring
growth releases in rings of trees that survived the outbreak. The rings of this tree provide evi-
dence of an outbreak in 1942.
1700 1760 1800 1850 1900 1950 2000
2.0
1.5
1.0
0.5
0.0
Mean
Doubling of the mean gr owth rate
SOURCE: ED BERG AND TOM VEBLEN

for example, suggest that spruce beetle outbreaks occurred frequently over the
last 250 years, with a mean return interval of 52 years. Similar patterns were
observed for spruce beetles in Colorado spruce forests and for mountain pine bee-
tles in lodgepole pine forests of British Columbia.
Are the Current Outbreaks Different?
The history of bark beetle infestations varies from forest to forest and ecosystem
to ecosystem, making generalization difficult. Research into past outbreaks, how-
e v e r, suggests that the current infestations may be more severe and widespread
than previous outbreaks due to a number of factors.
Regional scale: Although large outbreaks previously occurred in forested sys-
tems across the West, the historical evidence suggests that in the past century or
two, there have not been so many concurrent outbreaks at such a broad scale
involving so many species and locations simultaneously. This is particularly true
for the mountain pine beetle and spruce beetle. Similar widespread outbreaks
may have occurred farther back in geologic time, but we do not have the tools to
precisely evaluate the extent of outbreaks that occurred so far in the past.
Longer duration and intensity: In several ecosystems, outbreaks are lasting
longer than they typically did during the 20th century, and more trees are dying
at a faster rate. We are seeing massive tree mortality in habitats where previ-
ously recorded outbreaks have been limited in duration and extent.
Extended range: Most bark beetles are limited in geographical spread to the
range of host trees they attack. The spruce beetle for instance, can survive in all
places where spruce trees currently exist. The mountain pine beetle's range,
however, is bounded not by the trees it infests, but by climate. Lodgepole pine
extends farther north than the traditional range of the mountain pine beetle that
feeds on it, and ponderosa and other pine species grow farther south. Recently,
BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
20
Features of Current Bark Beetle Outbreaks
Current outbreaks appear to be different from observations of 20th
century historical outbreaks because of these factors.
• Regional scale – simultaneous severe outbreaks of several species are
occurring across the West
• Longer duration and intensity – some outbreaks are lasting longer and
more trees are dying at a faster rate
• Extended range – mountain pine beetles are moving north and east
beyond their previously recorded range
• New tree species – mountain pine beetles have been found
reproducing in spruce and jack/lodgepole pine hybrids

21
however, mountain pine beetles have dis-
persed out of their historical range, extending
farther north and east across Canada’s Rocky
Mountain crest (Figure 7). Currently, their
range has not expanded to the south, despite
the presence of suitable host trees.
New tree species: Although jack pine is not
believed to be a historical host for mountain
pine beetle, the insects have recently been
found in jack pine/lodgepole hybrids on the
east side of the Continental Divide in north-
ern Alberta, Canada. In addition, mountain
pine beetle has been found attacking and suc-
cessfully reproducing in spruce trees in
British Columbia and Colorado—the first evi-
dence of multiple generations of mountain
pine beetle in a non-pine species.
FIGURE 7.
Mountain Pine Beetle Range Expansion
SOURCE: NATURAL RESOURCES CANADA, CANADIAN FOREST SER VICE; USDA FOREST
SERVICE, FOREST HEALTH PROTECTION
The range of the mountain pine beetle generally follows its major
host pine species throughout western North America (shown in
green). The mountain pine beetle’s current range extends south to
the tip of southern California and central Arizona near Flagstaff. In
recent years, mountain pine beetle outbreak populations have been
found farther north in British Columbia and east in northern Alberta
than was observed in historical records, including those of the most
recent large outbreak in 1985.

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
22
South-Central Alaska — A Spruce Beetle Outbreak That Ran Its Course
From 1987 to 1997, an unprecedented run of warm summers, combined with mature, dense
forests, allowed spruce beetle populations in south-central Alaska to explode. By the end of
that 10-year period, the spruce beetle had essentially exhausted the supply of large host trees.
On the Kenai Peninsula, 1.6 million acres of forest were affected; greater south-central Alaska
lost most of the mature trees across a 2.9-million-acre area.
Weather in the region is strongly influenced by the El Niño-La Niña cycle—when fluctuating
Pacific ocean temperatures alter weather patterns across the western hemisphere. In the past,
several warm El Niño summers in a row could initiate an outbreak, which would then subside
with the onset of several cool La Niña summers. Following 1987, however, summer
temperatures increased about 2 °C on average. Temperatures did not drop below the mean,
even during the weak La Niña summers of 1988-89 and 1996. Warm summers accelerated the
spruce beetle’s normal two-year life cycle to one year, providing a double dose of beetles the
following spring. The warm weather also lengthened the growing season for host trees.
Without a concomitant increase in precipitation, however, the trees became stressed and more
susceptible to successful bark beetle colonization.
In addition to the unusually warm weather, the Kenai forests were especially susceptible
because they were densely stocked with large, mature trees. With continuing warm
temperatures enhancing spruce beetle development and survival, the only factor limiting the
outbreak was the supply of suitable host trees in the area. Only when the supply of host trees
had been exhausted did the outbreak collapse.
Will the overwhelming supply of dead trees in this large area of beetle-killed forests lead to
additional or more intense forest fires in the area? Soil charcoal analyses suggest that the
mean fire return interval in these forests is 400-600 years, while bark beetle outbreaks occur on
average every 50 years, according to tree-ring evidence. Therefore, there is no evidence of any
relationship
between spruce
beetle outbreaks
and fire over the
last 250+ years. The
climate during this
period was
distinctly cooler,
however, and
today’s warmer
climate may
intensify fire risk in
both green and
beetle-killed
forests.
Spruce beetle outbreaks in south-central Alaska have essentially exhausted the supply of
large host trees. On the Kenai Peninsula, 1.6 million acres have been affected.

W
hy are the recent outbreaks so severe and widespread? Scientists point to a
number of factors contributing to the outbreaks. Many of these factors,
such as rising temperatures, regionwide drought, and mature and over-mature
forests, are in effect across large areas in numerous ecosystems. Specific condi-
tions within particular forests or forest stands, however—such as the species and
density of trees, along with local ecology and weather conditions—often preclude
making broad generalizations.
Scientists do agree that the current outbreaks are taking place at a time when
forests have been affected by a variety of human activities, and that certain basic
dynamics appear to lie behind the current outbreaks. These include:
• a changing climate affecting all areas with bark beetle outbreaks;
• previous forest management practices such as selective timber harvest-
ing and wildfire suppression in some forest types and some geographic
areas;
• natural disturbances, such as climate-driven wildfire, that occurred in
previous centuries; and
• other human influences on forest ecosystems such as pollution.
Climate
Because bark beetles are highly sensitive to slight changes in their environ-
ments, small shifts in one component of their ecology—such as climate—can cre-
ate rapid and extreme shifts in outbreak dynamics. In recent years, the climate
in which bark beetles develop has changed noticeably. The 11 years between
1995 and 2006, for instance, ranked among the warmest since record-keeping
began in 1850; the current prolonged drought across the West is the longest in
duration since at least 1900; atmospheric carbon dioxide has increased 30 per-
cent over the last 150 years. All of these changes appear to have dramatic direct
and indirect effects on beetle populations.
•Direct Effects of Warming Temperature: The increase in regional
temperatures in recent years has permitted some species of bark beetle
to extend their ranges to higher altitudes and latitudes. Warming tem-
peratures in winter, fall, and spring also reduce cold-induced bark bee-
tle mortality. Research suggests that the probability of mountain pine
beetle survival during the past 10 to 20 years has increased as a result
of elevated minimum temperatures at many locations.
In addition, warmer temperatures have sped up the life cycles of some
species of bark beetles, increasing the number of beetles, which can
then more easily overcome a tree’s natural defenses. In some high-eleva-
tion forests, spruce beetles, which typically produce one generation
23
Contributing Factors

Climate Change and Bark Beetles: The Perfect Storm
The West’s changing climate—rising temperatures and decreasing
precipitation—has created weather conditions that are ideal for bark beetle
outbreaks.
• Because bark beetles are extremely sensitive to changes in
temperature, recent rising temperatures have led to rapid population
increases in some forests. Longer, warmer summers have extended
reproductive and growth periods, while fewer cold snaps and higher
winter temperatures have permitted increased bark beetle survival in
winter, spring, and fall.
• Because most tree species are sensitive to water stress, warm regionwide
droughts—such as the current prolonged drought across the West, the
longest we have
seen for over a
century—weaken
trees and make
them more
susceptible to bark
beetle attacks.
Forests full of
drought-stressed
trees, combined
with rapidly
expanding bark
beetle populations,
can combine to fuel
exponential beetle
population growth.
Mean Annual Temperature Trend
in the Western U.S.
1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010
Year
53
52
51
50
49
48
47
SOURCE: KELLY REDMOND, WESTERN REGIONAL CLIMATE CENTER
Annual January-December temperature in 11 western
U.S. states. Blue line is an 11-year running average.
every two years, have shifted to one generation per year. Some high-ele-
vation mountain pine beetles, as well, now produce a new generation in
one year instead of two. In the southwestern U.S., it is believed that
Ips
species, fueled by higher temperatures, also have been reproducing at a
faster rate. These population increases, in conjunction with drought,
have caused massive piñon and ponderosa mortality in the Southwest.
In Arizona, for instance, more than 6 million trees were killed across 1.2
million acres between 2001 and 2004.
•Indirect Effects of Warming Temperature and Drought: The cur-
rent regionwide drought, in combination with elevated temperatures,
has weakened trees throughout western North American forests.
Although moisture stress can kill trees directly, severely moisture-
stressed trees also may be more vulnerable to bark beetle attacks. The
BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
24
Mature forests
are the loaded
gun for today’s
severe bark
beetle
infestations,
and weather is
the trigger.

25
Piñon Ips in the Southwestern U.S.:
Drought, Overgrowth, and Rising Temperatures
In the southwestern United States, the interaction
between severe water stress and bark beetles has killed
ponderosa and piñon pines on millions of acres since
2000. Although bark beetle outbreaks also occurred
during the last severe drought in the Southwest region
in the 1950s, piñon pine mortality has been particularly
extreme during the recent drought.
Scientists believe there are two reasons for the current
widespread dieback. First, southwestern forest stands
are denser than they were 50
years ago. Historical livestock
grazing and fire suppression, along
with an unusually wet period in
the Southwest from 1978 until the
mid-1990s, promoted tree
establishment and resulted in
unusually dense forests and
woodlands. Overcrowded, stressed
trees are more vulnerable to bark
beetle infestation, especially
during dry conditions.
Second, although temperature
patterns were not unusual during
the 1950s drought, the recent
2000s drought has been
associated with significantly higher
temperatures. These warmer
temperatures amplified water
stress, in some cases causing
direct piñon pine mortality, and
also may have allowed
Ips
bark beetles to reproduce at a faster rate, increasing the number of
generations that develop in a single year. The result is unprecedented tree mortality across
extensive areas—95 percent of mature piñon pines have been killed in some forests of the
southwestern U.S., with over 3 million acres affected by substantial piñon pine mortality.
This recent drought episode is an example of “global-change-type drought,” where reduced
precipitation, accompanied by increased temperatures, can result in extensive and rapid
changes in vegetation. Climate models suggest that we will see more of this type of drought in
the southwestern United States in coming years.
PHOTO BY CRAIG D. ALLEN
Piñon pine mortality in the Jemez Mts. near
Los Alamos, NM, October 2002.
Piñon and Ponderosa Pine Mortality in
the Southwestern U.S., 1996-2005
Ips
and western pine beetle affected more than 7 million acres of
piñon and ponderosa pine in Colorado, Arizona, New Mexico, Nevada,
and Utah between 1996 and 2005. Severe drought and increased tem-
perature contributed to the widespread tree mortality.
1996 1997 1998 1999 2000 2001 2002 2003 2004 2005
Year
450,000
400,000
350,00
300,000
250,000
200,000
150,000
100,000
50,000
0
SOURCE: USDA FOREST SERVICE, FOREST HEALTH PROTECTION

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
relationship between moisture
stress and tree defense against bark
beetle attack, however, is complex.
Not every drought triggers a bark
beetle outbreak, and different forest
ecosystems respond differently to
outbreaks triggered by moisture
stress. In the southwestern piñon
and ponderosa pines affected by the
recent
Ips
and western pine beetle
outbreaks, the return to more typi-
cal moisture levels also meant the
end of the beetle outbreak. In the
ongoing mountain pine beetle out-
break in British Columbia, however,
beetle population growth and associated tree mortality continued even
after drought conditions subsided.
Disturbance and Human Influence
•Wildfire Suppression:
Since the early 20th century when the U.S. and
Canadian governments implemented a policy to suppress all fires on federal
land, many fire-prone ecosystems have experienced long fire-free intervals.
In some cases, the species composition and structure of those forests have
changed, creating dense forests full of the mature and over-mature trees
bark beetles favor.
Such shifts have taken place in low-elevation ponderosa pine forests in the
southwestern United States, where once-parklike stands of trees, kept open
by regular surface fires every two to 12 years, are now dense forests, and in
lodgepole pine forests in British Columbia. Other ecosystems, however, such
as subalpine spruce, fir, and lodgepole pine forests in Colorado, and white-
bark pine forests in the northern Rocky Mountains, typically experience
longer periods of time between fires. In many areas, current forest density
does not appear to be a consequence of fire suppression over the last century.
The role of fire suppression in bark beetle outbreak dynamics is a topic of
much discussion among scientists, reflecting the need for additional
research. Although the effect of fire suppression on bark beetle outbreaks
varies by forest ecosystem, region, and the level of management applied, it is
fair to conclude that in some cases, fire suppression policies may have helped
create a landscape that is more susceptible to bark beetle attacks.
•Disturbance history: Bark beetle-caused tree mortality can be signifi-
cant in forests where the majority of trees are the same species and are
26
New Inputs
Scientists believe a number of
factors have changed the
“inputs” to forest systems in
which bark beetles are found,
and have contributed to recent
outbreaks. They include:
• Rising temperatures
• Prolonged drought
• Air pollution
• Previous forest
management practices

uniformly mature and large. Because aggressive bark beetles favor
mature trees, stands that have regrown or been replanted after a dis-
tinct disturbance event, such as harvesting, wildfire, or previous bark
beetle outbreaks, may be more vulnerable to future bark beetle out-
breaks, especially when the majority of trees are competing for
resources in overcrowded conditions. For instance, in British Columbia
selective harvesting—along with fire suppression—has contributed to a
predominance of mature stands that are highly susceptible to mountain
pine beetle outbreaks (Figure 10). Not all disturbances create suscepti-
ble forests, however. Some fires, and even
some timber harvesting, can reduce stand
density and future susceptibility to bark-bee-
tle outbreak initiation.
•Air Pollution: Local and long-distance disper-
sal of air pollution from heavily populated
areas and increasing development on the
edges of forests also can have an indirect
influence on bark beetle outbreaks. Ozone can
damage needles, disrupting the tree’s phot-
synthetic capacity, thereby weakening the
tree and making it more susceptible to bark
beetle attack. An increase in atmospheric
nitrogen deposition can stimulate growth in
trees, leaving energy resources too depleted to produce sufficient resin
to defend against bark beetle attack.
27
Are the current bark beetle
outbreaks unprecedented?
Relative to what we know about the scale
of historic outbreaks, many of the current
bark beetle outbreaks do appear to be
larger, more widespread, more severe,
and occurring in new and novel habitats.
This is, in part, due the fact that the
inputs to the system have changed,
allowing bark beetles to thrive.
FIGURE 8.
Susceptible Lodgepole Pine Forests in British Columbia
40 80 120 160 200 240
1910 1930 1950 19901970
17% 26% 35% 49%
40 80 120 160 200 240 40 80 120 160 200 240 40 80 120 160 200 240 40 80 120 160 200 240
5
4
3
2
1
0
Forest Age Class (years)
53%
SOURCE: TAYLOR AND CARROLL 2004.
In British Columbia, the percentage of lodgepole pine forest in age classes susceptible to mountain pine beetle (in red) is
predicted to have increased from 17 percent of total forest area in 1910 to 53 percent in 1990. A century of fire suppression
and forest management practices have helped create these forest conditions.

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
W
hile some bark beetle researchers comb through tree-ring, pollen, and fossil
records to understand how and why current beetle outbreaks may differ
from those in the past, others scientists have developed sophisticated simulation
models to help predict bark beetle activity in the near and distant future.
These models use data gathered in the field and laboratory to describe the
mechanics of bark beetle response to temperature. Such mechanistic models can
then be used to predict how current and future temperature patterns may affect
bark beetle outbreak dynamics on local and landscape scales (Figure 9). These
temperature-based models have been developed only for the mountain pine bee-
tle and spruce beetle. They do not yet predict dynamics of other aggressive bark
beetle species, nor do they take into account variability in temperature response
among mountain pine and spruce beetle populations across their ranges. To ade-
quately predict the effect of changing climate on all forest ecosystems throughout
western North America, scientists must develop models for other species, and
refine the current models to take into account geographic variability within a
species.
There is always a high degree of uncertainty in attempting to forecast the future,
especially on a regional or global scale. Still, scientists can make informed esti-
mates by using these sophisticated computer models, combined with an under-
standing of the underlying mechanisms of bark beetle response to temperature,
to predict how those dynamics may adjust to changing conditions. The general
consensus is that continued warming will fuel beetle attacks in areas where bee-
tle activity was previously constrained by climate, such as in northern latitudes
and at higher elevations. In locations where bark beetles are currently success-
ful, however, continued warming could cause some beetle populations to go local-
ly extinct unless they are able to rapidly adapt. If local populations of bark bee-
tle die out due to warming, it is possible that other species could then move into
those vacated niches. For example, aggressive species currently restricted to the
southwest U.S. and Mexico could expand in range northward as the climate
warms.
Ecological Consequences of Recent Bark Beetle Outbreaks
Bark beetles have, for millennia, been a natural part of the forest regeneration
process. Bark beetles help to winnow out old and mature trees so that younger,
more productive trees can replenish an aging forest. They also accelerate the
process of tree decay to help forests capture and recycle nutrients. In recent
years, however, a combination of factors, including warm temperatures, drought-
stressed trees, susceptible landscapes, and historical management practices, may
have tipped many systems out of the balance we have observed over the past
century. Beyond the troubling sight of vast areas of dead trees scattered over
large landscapes in western North American forests, scientists are concerned
that the current levels and rates of tree mortality in some forest ecosystems may
28
The Future of Our Forest Ecosystems

be pushing these systems beyond their ability
to recover and regenerate.
Bark beetles can kill large numbers of trees in
a forest within three to five years, yet many
tree species require hundreds of years to reach
maturity. Therefore, a massive beetle outbreak
can create large shifts in vegetation types and
patterns in affected forests. In a stand with
several different species of trees, those tree
species that are not attacked may now become
the dominant vegetation—for instance, aspens
or fir trees may take over after spruce trees
die.
In stands that are composed predominantly
of one species of similar age, a bark beetle
outbreak can cause an even more dramatic
shift in the type of vegetation found there.
29
FIGURE 9.
Predicted change in Bark Beetle Outbreak Probability with Warming
Temperatures
Temperature-driven model predictions suggest that over the next 30 years, spruce forests in Alaska, the Northwest
Territories, and at high elevations in the western United States will see increased probability of univoltine life cycle spruce
beetle. Higher temperatures are expected to speed up the life cycle of many spruce beetles from one generation every two
years to one generation per year, resulting in exponential population growth and thus an increased likelihood of severe out-
breaks (left). Over the same time period, the likelihood of population success in areas within the current range of mountain
pine beetle (see Figure 7) is predicted not to change dramatically except in high elevation forests of the western U.S. and in
northern British Columbia (right). Over the next 30 years, a low likelihood of mountain pine beetle range expansion is pre-
dicted in central and eastern U.S. and Canada, areas where pine species grow but are outside the current range of mountain
pine beetle.
Grass invading areas on Alaska’s Kenai penin-
sula, where overstory spruce were killed by
spruce beetle and subsequently removed by
logging.
PHOTO BY ED BERG
1.0
0.0
-1.0
Change in
Probability

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
30
Whitebark Pine in the Northern Rocky Mountains: Mountain
Pine Beetle Outbreaks, Coupled with the Exotic Blister Rust Pathogen,
Threaten Local Extinction
In the past 10 years, mountain pine beetles have infested and killed more than 570,000 acres
of whitebark pine in the northern Rocky Mountains of the United States. Whitebark pine is a
long-lived five-needle white pine that lives in the upper subalpine zone, tolerating harsh
temperature and moisture extremes that many other conifers are not adapted to. Whitebark
pine trees are vital to the survival of a number of wildlife species, such as red squirrels, black
and brown bears, and Clark’s nutcracker—the tree’s principal seed disperser. Whitebark pine
trees also can act as "nurse trees" for other species such as subalpine fir by providing shelter
from the harsh, windy conditions. The trees grow higher in elevation than most conifers, and
both stabilize soil and regulate snow melt in the treeline zone.
Although bark beetle outbreaks have occurred infrequently in whitebark pine forests in the
past century, written, tree-ring, and fossil records indicate that during the late 1920s and
1930s, elevated temperatures and reduced precipitation were associated with widespread
mountain pine beetle-caused whitebark pine mortality throughout the northern Rocky
Mountains. Once temperatures cooled, the infestations subsided. With projected climate trends
suggesting a long run of warm temperatures, however, today’s severe beetle outbreaks show
no sign of letting up.
Mountain Pine Beetle-Caused Mortality
in High Elevation White Pines
SOURCE: USDA FOREST SERVICE, FOREST HEALTH PROTECTION
More than 1.3 million acres of high elevation
white pines (including whitebark pine, limber
pine, and bristlecone pine) in the western
United States have been affected by mountain
pine beetle over the past 10 years. California is
not included in these figures.
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006
Year
300
250
200
150
100
50
0

31
Mountain pine beetle-killed whitebark pine in Yellowstone National Park.
PHOTO BY JEFF HICKE
These mountain pine beetle outbreaks come at a time when a non-native
disease, white pine blister rust, has infected more than 65 percent of
whitebark pine stands in the northern Rocky Mountains. Blister rust can
reduce cone production, thereby limiting the number of new seeds available
to renew those forests. When the infection is severe, the affected trees die
as well. The combination of blister rust infection and bark beetle infestation
is particularly troubling because bark beetles are killing trees that have
developed genetic resistance to blister rust. The combination of blister rust
and mountain pine beetle thwarts the potential benefits of natural selection.
Resistant trees survive blister rust, but then succumb to mountain pine
beetle. This one-two punch could mean the end of whitebark pine in parts of
its range where both mountain pine beetles and blister rust are currently at high levels.
With so many trees dying, land managers and ecologists are developing restoration programs to
prevent local and regional extinction of whitebark pine forests. Although it is impossible to suppress
mountain pine beetle populations once they have become large enough to successfully colonize large
numbers of trees in a stand, the direct, manual application of pesticides can protect individual, high-
value trees from mountain pine beetle attack. Widespread pesticide application is not feasible over
large areas, however, especially at the remote, high elevation sites where
whitebark pines are found. Synthetically produced pheromones, which
mimic compounds naturally produced by the beetles, are easier to apply
and can provide some protection against fatal bark beetle attacks, but
have not proven consistently effective over several years.
Forestry professionals are also looking at long-term restoration efforts to
develop whitebark pine communities that can weather the dual threats of
blister rust and mountain pine beetle. Strategies include planting rust-
resistant seedlings in areas where losses have been high, as well as
stimulating natural regeneration of the stands. Allowing surface fires to
burn in some affected whitebark pine stands can reduce competition with
other vegetation and promote better growing conditions for those trees
that survive the current outbreaks.
The Clark’s nutcracker is one
of a number of species that
depend on the whitebark
pine for survival.
PHOTO COURTESY OF
WHITEBARKPINEFOUND.ORG
Blister rust is an exotic pathogen
that infects five-needle pines.

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
On the Kenai peninsula in Alaska, for instance, where
spruce beetle recently killed millions of trees over a rela-
tively short period of time, native bluejoint grass has
thrived in the now-open canopies. This ingrowth of grasses
has been most pronounced in areas that were salvage-
logged following the outbreak. Spruce trees may not be able
to grow on many of these sites for hundreds of years,
because the grass crowds out spruce seedlings as they
regenerate. Outbreak areas that were not logged, however,
sustained enough “nurse trees”—fallen trees that provide
shelter for new seedlings—to foster a new cohort of spruce
and other tree species.
Such rapid changes in forest composition also can affect
wildlife species that have evolved to survive in specific habi-
tats. Although some bird species rely on bark beetle-infested
trees to nest and feed, other species depend on live trees. The
Clark’s nutcracker, for instance, feeds on whitebark pine
seeds, which the birds hide away in thousands of caches to
sustain them during lean times. Red squirrels also store large numbers of white-
bark pine cones in middens on the forest floor, and grizzly and black bears regu-
larly raid those middens to feed during the critical months prior to winter hiber-
nation. Similarly, piñon pine seeds are vital food sources for many native mam-
mals and birds in the southwestern U.S. When piñon pines experience mass
mortality, the species that depend on them may decline as well.
In addition, massive tree mortality can affect watershed quality and quantity.
Live trees in high-elevation watersheds provide shade and shelter that help to
maintain the winter snowpack and prevent quick runoff during the spring melt
and summer storms. Large numbers of bark beetle-killed trees within a water-
shed increase the risk of rapid snow loss and can enhance annual streamflow.
Live trees also provide aesthetic and recreational forest values, and wilderness
tourism may decline as a consequence of massive bark beetle-kill events. A mas-
sive beetle kill on the slopes of ski resorts, for example, can result in earlier
melting in spring and more drifting during wind events, compromising snow
quality. In addition, bark beetle-killed trees may become “hazard trees” because
once dead they are prone to falling, and require costly removal to keep backcoun-
try trail networks and forest campgrounds safe and navigable.
Bark beetle outbreaks also can modify forest carbon balances. Mature forests
store large amounts of carbon. When mature trees are killed by bark beetles, the
stored carbon in the dead trees is released over several decades during decompo-
sition. Thus, in the short term, forest carbon storage can be greatly reduced as
32
Bark-beetle-caused tree
mortality surrounding
homes near Lake
Arrowhead in southern
California.
PHOTO BY LAURA MERRILL

the stored carbon is released. However, as new trees
grow in the several decades following an outbreak,
the net carbon flux (a measure of both decomposi-
tion and new growth) eventually shifts, and the for-
est again becomes a carbon sink, absorbing more
carbon than it releases. An adequate estimate of the
influence of bark beetle outbreaks on carbon bal-
ance requires consideration of multiple factors,
including regeneration rates, the amount of time
since the outbreak occurred, and the number and
vigor of surviving trees. More research is needed to
assess how changes in outbreak regimes might
affect ecosystem carbon balances.
Bark Beetles and Fire
Bark beetles and fire are the two biggest natural
disturbances in western North
American forests, often—but not
always—interacting to regenerate old
stands. Fire ecologists are just begin-
ning to understand the relationship
between bark beetle-caused tree mor-
tality and fire in those ecosystems
they have studied. This relationship is
extremely complex, and varies by loca-
tion and forest type. Scientists use the
term “fire hazard” to describe the
probability of a fire occurring in a for-
est. “Fire hazard” refers to the state of
the fuels in a given stand—the density
of the overhead canopy and the
amount and arrangement of ground
and ladder fuels including grasses,
shrubs, and small trees. Weather also
influences fire behavior, but the term
“fire hazard” refers specifically to the
state of fuels independent of those
variables such as temperature, wind,
and precipitation that influence fuel
moisture content.
Figure 10 is a generalized framework
summarizing current knowledge of fire
33
Crown-fire hazard remains high in the
year or two after a bark beetle out-
break, when dead needles remain on
trees. After the needles drop, crown-
fire hazard also falls.
FIGURE 10.
Hypothesized Fire Hazard Following
Bark Beetle Outbreaks
SOURCE: CRAIG ALLEN
Hypothesized changes in conifer forest fire hazard following a severe bark
beetle outbreak assuming continuously dry climate conditions. Actual fire
behavior will vary depending on many factors, including predominant cli-
mate and fire weather, landscape site factors (e.g., slope), and vegetation
structure and species. We refer the reader to publications in the Additional
Reading section for more detail on these complex relationships.
TIME
HIGH
LOW
Extensive tree
mortality.
Dead needles
still on trees.
Canopy Fire
Surface Fire
Live, water-
stressed
conifer
forest.
Dead needles drop.
Fine surface fuels
increase. Surface
fuels drier.
Dead trees start to fall.
Herbs, shrubs,
and trees regrow.
Coarse woody surface
fuels increase.

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
hazard and fire behavior following
a bark beetle outbreak and is based
on field research and simulation
model results. Both crown- and
surface-fire hazard change through
time following a bark beetle out-
break event in a stand of living
conifers. For one to two years fol-
lowing a bark beetle outbreak,
crown-fire hazard is expected to be
high while dead needles remain on
the tree, stocking the canopy with
dry, fine fuels that can ignite quick-
ly during weather conditions con-
ducive to fire. After the dead nee-
dles fall and canopy fuel continuity is interrupted, crown-fire hazard declines
markedly relative to pre-outbreak conditions. However, surface-fire hazard
may increase in that period. Wind speed and solar input increase without
the shade and shelter provided by the high canopy, drying out fine surface
fuels such as the fallen needles from beetle-killed trees. Warmer, windier
conditions also allow fires to spread more quickly and burn more intensely.
Surface-fire hazard continues to increase once the dead trees fall, adding
downed wood to the surface fuel load. Shrubs and young trees that grow in
place of the bark beetle-killed trees also add to surface-fire
hazard and serve as ladder fuels that can spread fire into
tree crowns.
The effects of bark beetle outbreaks on subsequent fire
hazard varies greatly depending on ecosystem type and ini-
tial stand conditions (e.g., understory and overstory compo-
sition, stand structure and age, and the number of stand-
ing and fallen dead trees). Generally speaking, crown- and
surface-fire risks change with time following outbreaks,
and factors such as weather and forest composition play
large roles in determining whether and how intensely a
fire will burn. Results from two studies illustrate these
points.
Research into the effects of a spruce beetle outbreak in
Colorado during the 1940s indicates that fire frequency
was no greater in the half-century following the outbreak
than in the years preceding it when compared with forests
not affected by the spruce beetle outbreak (Figure 11).
Widespread fires occur only an average of once every two
34
Crown fires spread rapidly and can destroy large areas
of forest.
FIGURE 11.
Fires in Colorado Following
Spruce Beetle Outbreaks
SOURCE: BEBI ET AL 2003
A large spruce beetle outbreak in the White River
National Forest in Colorado during the 1940s had
little influence on the number and size of subse-
quent fires.
1940s Spruce No Outbreak
Beetle Outbreak
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0

centuries in high-elevation spruce forests, and are generally depend-
ent on extreme drought. Despite an increase in fire hazard following
the 1940s outbreak, weather conditions were generally not conducive
to extreme fire events during those years. Only during the severe
drought of 2002 did those forests burn. When they did, both beetle-
killed stands and stands unaffected by the spruce beetle burned in a
similar fashion. These observations suggest that many of Colorado’s
high-elevation spruce-fir forests have burned not because of a large
change in fuels following a spruce beetle outbreak, but because of
“fire weather”— hot, dry, and windy conditions that promote acceler-
ation of severe crown fires.
Bark beetle outbreaks appear to have a more causal relationship with fire in
other ecosystems, such as lodgepole pine forests in Yellowstone National
Park. In the extensive Yellowstone fires of 1988, lodgepole pine forests that
had been affected by a mountain pine beetle outbreak 15 years previous
were more likely to burn than were nearby locations where bark beetles had
not been active. Trees killed by an outbreak that had occurred only five to
ten years previously, however, were not found to have contributed to the
fire’s spread or increased its intensity. This evidence indicates that, in the
areas studied in Yellowstone, the release of understory “ladder fuels” years
35
The Yellowstone fires burned intensely both in areas where beetles had been active and in are a s
w h e re they had not. Studies indicate that trees were slightly more likely to burn in areas where
beetles had been active in an outbreak that occurred 15 years before the fires. However, an
a rea that experienced an outbreak 5-10 years previous to the fires was less likely to burn .
With Bark Beetles,
One Size Does Not
Fit All
The factors influencing a
bark beetle outbreak
vary depending on the
species of bark beetle,
geographic region, and
host tree species.

BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES
after a mass beetle-kill poses more of a fire risk than the short-term increase
in dead fuel caused by a more recent outbreak. Although beetle-affected
stands played varying roles in the Yellowstone fires, the principal reason
why the fires were so intense was that weather conditions were extreme,
with prolonged drought and high winds. Indeed, some forests without previ-
ous bark beetle activity burned as intensely as forests where beetles had
been especially active.
If bark beetle outbreaks sometimes have an effect on forest fire hazard, the
reverse is also true: forest fires can influence bark beetle activity. A l t h o u g h
stand-replacing fires can reduce outbreak risk in the long-term by removing the
large, mature trees that bark beetles prefer, in the short term, some species of
bark beetles, in particular Douglas-fir beetle and western pine beetle, are
attracted to trees that have been damaged, but not killed, by fire. Post-fire bee-
tle attack by these species can kill up to 25 percent of trees that survive a forest
fire—a factor that may need to be taken into account when considering the use
of prescribed burns to promote forest health. We do not currently have sufficient
information, however, to assess whether fire-injured trees can help foster the
build-up of bark beetle populations to such an extent that they can attack and
overcome healthy trees in the surrounding forests.
These studies help us understand the historical relationship between bark bee-
tles and fire. As bark beetle outbreak dynamics shift—and we see levels of tree
mortality that have not been observed in recent history—the dynamic relation-
ship between fire and bark beetle disturbances also may change. Although beetle
outbreaks and fire risk differ from forest to forest, historical fire and beetle activ-
ity are certainly linked in one respect: conditions that influence wildfires—warm
temperatures, drought-stressed trees, and a dense forest canopy—also favor bee-
tle activity.
Research Gaps
Although outbreaks in recent years have provided scientists with excellent
opportunities to conduct studies and gather new information about the role of
bark beetles in western forests, much research remains to be done. In addition to
the gaps in our understanding discussed above, a number of bark beetle-related
dynamics are still not fully understood.
• How different species of trees are affected by climate change, including
seasonal shifts in precipitation, temperature, and increased atmospheric
gases, and the indirect effect of these changes on tree resistance to
attacking bark beetles.
• The degree to which the current beetle population explosions have been
triggered by changes in climate such as drought and rising tempera-
tures, versus forest habitat conditions such as overly dense or homoge-
36

37
nous stands of trees. Is it possible that shifts in climate may create cer-
tain forest habitat conditions that constrain bark beetle outbreaks?
• The impact of outbreaks on ecosystem processes such as forest fire,
nutrient cycling, and forest regeneration, and how current changes in
bark beetle outbreaks will affect these processes.
• How these dramatic new outbreaks will affect various landscapes, par-
ticularly those that have not previously, or have only rarely, experienced
bark beetle outbreaks in recorded history.
• Whether various harvesting strategies, including thinning at large spa-
tial scales prior to a beetle outbreak, reduce the probability and severity
of an infestation in multiple forest types.
• The interplay between beetle outbreaks and invasion by non-native
species. Do invasive plants such as cheatgrass, which has expanded into
areas of widespread piñon mortality in the southwestern United States,
thrive in areas that have been hard-hit by bark beetles?
• The economic and social impacts of bark beetle disturbance. How will
large-scale outbreaks and massive tree mortality affect the forest-prod-
ucts and tourism industries and the communities that depend on them?
• How does the loss of habitat associated with bark beetle outbreaks
affect wildlife—in particular threatened, sensitive, and endangered
species?

38
Bark Beetles and Forest Management
The current bark beetle outbreaks, and the vast numbers of dead trees these outbreaks leave behind,
p resent difficult challenges for managers and policymakers. Multiple factors—such as stand type, local
conditions, bark beetle population level, and re s o u rce and community objectives—must be considere d
b e f o re making landscape-scale management decisions in forests affected or threatened by bark beetle
infestations.
For example, it may make sense to use insecticides on high-value trees on a small scale, such as in
c a m p g rounds or around homes, but such spraying is not feasible on a large landscape scale. In some
instances, strategies such as thinning may be effective prior to a large outbreak to reduce competition
among trees, thereby enhancing the health and defensive capacity of individual trees. This strategy,
h o w e v e r, is less effective once bark beetle populations have grown so large that they can overcome the
defenses of even the healthiest trees. In addition, populations of certain bark beetle species, such as
those within the genus
I p s
, can in some instances re p roduce and proliferate in logging debris that
thinning operations leave behind. More o v e r, some environmental triggers, such as unusually warm
w e a t h e r, may override all attempts to prevent a successful colonization event. Once an outbreak re a c h e s
landscape scale, no known management options are effective or practical for stopping the outbre a k .
Evaluating the appropriate management response to a bark beetle disturbance event requires
understanding not only the complexities of local and regional ecology—the species-specific cross-scale
interactions occurring within the forested landscape—but also the long-term influence of these
management actions on the surrounding landscape. For example, in the wildland-urban interface
where homes have been built near forest boundaries, land managers may experience pressure to
remove beetle-killed trees to reduce perceived fire risk and lessen the visual impact of so many dead
trees. Although removing dead trees and other fuel has been shown to reduce fire risk in the
immediate vicinity of a home and is advisable under many circumstances, the link between bark beetle
outbreaks and subsequent fire at the larger landscape scale is not fully understood. In addition, the
role mechanical thinning should play in fire reduction is unresolved and varies depending on the
location and type of forest. Although conventional harvesting usually targets the largest timber for
removal, recent research suggests that in some ecosystems, the removal of small trees, brush, and
ground fuels that carry a fire may actually be more effective in reducing fire risk.
The changing dynamics of current outbreaks make management decisions even more difficult. Some
outbreaks have no recent precedent, so an appropriate management response can not necessarily be
formulated based on previous events. One important aspect of future forest management will be an
evaluation of multiple approaches across a range of spatial scales and outbreak severity levels. Many
areas will regenerate naturally following a bark beetle outbreak and require no action. In some areas
severely affected by recent outbreaks, land managers may want to consider the creation of a diverse
forest through modifications to species and age classes at a regional scale. In addition, some
ecosystems that have highly susceptible forest conditions, but are currently unaffected by bark
beetles, may benefit from actions to reduce stand density. This is particularly true in lodgepole and
ponderosa pine stands where research has shown that thinning can reduce susceptibility. Finally, policy
makers and forestry professionals should incorporate projections of climate-change induced stress on
host forests and direct effects of warming temperature on bark beetle populations when developing
future forest management strategies.
BARK BEETLE OUTBREAKS
IN WESTERN NORTH
AMERICA: CAUSES AND
CONSEQUENCES

39
Additional Reading
Allen, C.D. 2007. Cross-scale interactions among forest dieback, fire, and erosion in
northern New Mexico landscapes. Ecosystems 10:797-808.
Aukema, B.H, A.L. Carroll, Y. Zheng, J. Zhu, K.F. Raffa, R.D. Moore, K. Stahl, and
S.W. Taylor. 2008. Movement of outbreak populations of mountain pine beetle:
Influences of spatiotemporal patterns and climate. Ecography 31:348 – 358.
Bebi, P., D. Kulakowski, and T.T. Veblen. 2003. Interactions between fire and spruce
beetle in a subalpine Rocky Mountain forest landscape. Ecology 84:362-371.
Bentz, B.J., J. A. Logan, and G.D. Amman. 1991. Temperature dependent develop-
ment of the mountain pine beetle (Coleoptera: Scolytidae), and simulation of its
phenology. The Canadian Entomologist 123:1083-1094.
Bentz, B.J., J.A. Logan, and J.C. Vandygriff. 2001. Latitudinal life history variation
in
Dendroctonus ponderosae
(Coleoptera: Scolytidae) development time and size.
The Canadian Entomologist 133:375-387.
Berg, E.E., J.D. Henry, C.L. Fastie, A.D. De Volder, and S.M. Matsuoka. 2006.
Spruce beetle outbreaks on the Kenai Peninsula, Alaska, and Kluane National
Park and Reserve, Yukon Territory: Relationship to summer temperatures and
regional differences in disturbance regimes. Forest Ecology and Management
227: 219–232.
Bigler, C., D. Kulakowski, and T. T. Veblen. 2005. Multiple disturbance interactions
and drought influence fire severity in Rocky Mountain subalpine forests. Ecology
86:30183029.
Breshears, D.D., O.B. Myers, C.W. Meyer, F.J. Barnes, C.B. Zou, C.D. Allen, N.G.
McDowell, and W.T. Pockman. (In press). Tree die-off in response to global-
change-type drought: Mortality insights from a decade of plant water potential
measurements. Frontiers in Ecology and the Environment. DOI:10.1890/080016.
Brunelle, A., G. Rehfeldt, B. Bentz, and S. Munson. 2008. Holocene records of
Dendroctonus
bark beetles in high elevation pine forests of Idaho and Montana,
USA. Forest Ecology and Management 255:836-846.
Cardoza, Y. J., K. D. Klepzig, and K. F. Raffa. 2006. Bacteria in oral secretions of an
endophytic insect inhibit antagonistic fungi. Ecological Entomology 31: 636-645.
Fettig, C.J., K.D. Klepzig, R.F. Billings, A.S. Munson, T.E. Nebeker, J.F. Negrón, and
J.T. Nowak. 2007. The effectiveness of vegetation management practices for pre-
vention and control of bark beetle infestations in coniferous forests of the west-
ern and southern United States. Forest Ecology and Management 238: 24-53.
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