Bark Beetle and Fire Interactions in Western Coniferous Forests: Research Findings

Page created by Dana Strickland
 
CONTINUE READING
Bark Beetle and Fire Interactions in Western Coniferous Forests: Research Findings
Bark Beetle and Fire
Interactions in Western
Coniferous Forests:
Research Findings
Christopher J. Fettig, Sharon M. Hood, Justin B. Runyon, and Chris M. Stalling

         ative bark beetles and wildfires     Figure 1—Tree mortality following a bark beetle outbreak in the Sierra Nevada in California.

N        are important disturbances
         in western coniferous forests.
Bark beetles can colonize and kill trees
                                              California experienced a severe drought from 2012 to 2015, stimulating a large bark beetle
                                              outbreak in the central and southern Sierra Nevada. Most tree mortality was caused by western
                                              pine beetle (Dendroctonus brevicomis), which readily colonizes drought-stressed ponderosa
of all species, ages, and sizes, but each     pine (Pinus ponderosa), but other tree and shrub species were also affected. About 89 percent
species exhibits unique host preferences      of the ponderosa pines in the three largest diameter classes were killed (Fettig and others 2019),
and impacts. Some bark beetles cause          representing the loss of an important structural component of these forests. Mortality of
extensive levels of tree mortality (table     sugar pine (Pinus lambertiana), caused primarily by mountain pine beetle (Dendroctonus
1), as demonstrated by mountain               ponderosae), was also substantial (48 percent). In total, 49 percent of the trees died between
pine beetle (Dendroctonus ponderosae)         2014 and 2017. Photo: C. Fettig, USDA Forest Service.
in several pines, western pine beetle
(Dendroctonus brevicomis) in ponderosa
pine (Pinus ponderosa), Douglas-fir beetle    In general, bark beetles require
(Dendroctonus pseudotsugae) in Douglas-       living phloem (the layer of cells
fir (Pseudotsuga menziesii), and spruce                                                           Christopher Fettig is a research entomologist
                                              within the inner bark that transports               for the Forest Service, Pacific Southwest
beetle (Dendroctonus rufipennis) in several   photosynthates (sugars) within the
spruces. Other bark beetles are secondary                                                         Research Station, Davis, CA; Sharon Hood is
                                              tree) to reproduce. When bark beetle                a research ecologist for the Forest Service, Rocky
agents that colonize stressed, dead,          populations are low, the beetles create
or dying trees. The impacts of these                                                              Mountain Research Station, Missoula, MT;
                                              small gaps in the forest canopy by                  Justin Runyon is a research entomologist for
secondary agents often go unnoticed,          colonizing and killing trees stressed by
while the former occasionally drive                                                               the Forest Service, Rocky Mountain Research
                                              age or other factors. During bark beetle            Station, Bozeman, MT; and Chris Stalling is a
headlines in large newspapers.                outbreaks, large numbers of trees can be            biologist for the Forest Service, Rocky Mountain
                                                                                                  Research Station, Missoula, MT.

Fire Management Today                                             14                                      JANUARY 2021 • VOL. 79 • NO. 1
Bark Beetle and Fire Interactions in Western Coniferous Forests: Research Findings
Table 1—Bark beetles recognized as causing substantial levels of tree mortality during outbreaks in the Western United States.
                                                                                                                      Current knowledge of effects on
         Common name                         Scientific name                     Common host(s)
                                                                                                                       wildfire behavior and severitya
                                                                         Lodgepole pine, sugar pine,
 California fivespined ips             Ips paraconfusus                                                                                Low
                                                                         ponderosa pine
 Douglas-fir beetle                    Dendroctonus pseudotsugae         Douglas-fir                                                   Moderate
                                                                         White fir, grand fir, California
 Fir engraver                          Scolytus ventralis                                                                              Low
                                                                         red fir
 Jeffrey pine beetle                   Dendroctonus jeffreyi             Jeffrey pine                                                  Low
                                                                         Whitebark pine, lodgepole pine,
 Mountain pine beetle                  Dendroctonus ponderosae           limber pine, sugar pine, western                              High
                                                                         white pine, ponderosa pine
 Northern spruce engraver              Ips perturbatus                   White spruce, Lutz spruce                                     Low
                                                                         Lodgepole pine, Jeffrey pine,
 Pine engraver                         Ips pini                                                                                        Low
                                                                         sugar pine, ponderosa pine
 Pinyon ips                            Ips confusus                      Pinyon pine(s)                                                Low
                                                                         Engelmann spruce, white
 Spruce beetle                         Dendroctonus rufipennis                                                                         Moderate
                                                                         spruce, Lutz spruce
 Western balsam bark beetle            Dryocoetes confusus               Subalpine fir                                                 Low
 Western pine beetle                   Dendroctonus brevicomis           Ponderosa pine, Coulter pine                                  Low

a. Level (low, moderate, or high) defined in relation to knowledge of the effects imposed by mountain pine beetle outbreaks, which have been most intensively studied.

killed over extensive areas (fig. 1), often               to repel beetle attack (Kolb and others                  In this article, we consider two common
adversely affecting timber and wood                       2016). Accordingly, recent bark beetle                   interactions between bark beetles and
fiber production, water quality and                       outbreaks have been correlated with                      wildland fires:
quantity, fish and wildlife populations,                  shifts in temperature and precipitation
opportunities for outdoor recreation,                     caused by climate change. In some                        1. The effects of fuel reduction
and biodiversity and carbon storage,                      forests, increases in tree density have                     treatments (prescribed fire and
among other ecological goods and                          exacerbated the effect by providing an                      mechanical thinning) and wildfires
services (Morris and others 2018).                        abundance of hosts and by increasing                        on bark beetles; and
                                                          competition among hosts for limited                      2. The effects of bark beetle outbreaks
HISTORIC OUTBREAK LEVELS                                  resources, making trees more vulnerable                     and associated levels of tree
The amount of tree mortality caused                       to beetle attacks.                                          mortality on fuels and wildfire
by bark beetles in the Western United                                                                                 behavior and severity.
                                                          Wildfires have sculpted many western
States has exceeded that caused by
                                                          forests for millennia, reducing the                      We briefly describe the current state
wildfires in the last 3 decades (Hicke
                                                          quantity and continuity of fuels,                        of knowledge and identify gaps in
and others 2016), and several recent
                                                          discouraging establishment of fire-                      knowledge needed to make informed
outbreaks are considered the most
                                                          intolerant tree species, and influencing                 management decisions.
severe in history. Since 2000, for
                                                          the susceptibility of forests to bark
example, about 25.5 million acres (10.3
                                                          beetle outbreaks and other disturbances.                 EFFECTS OF FUEL
million ha) in the Western United
                                                          Climate change is increasing the                         REDUCTION TREATMENTS
States have been affected by mountain
                                                          number of large wildfires (fires greater
pine beetle. Activity peaked in 2009,                                                                              ON BARK BEETLES
                                                          than 1,000 acres (400 ha) in size), the
with 8,842,698 acres (3,578,513 ha)                                                                                Tens of millions of acres of forest in the
                                                          frequency of wildfires, the length of
affected in that year alone.                                                                                       Western United States are classified as
                                                          the wildfire season (by up to 90 days
Bark beetles are cold-blooded                             in some locations), and the cumulative                   having moderate to high fire hazards.
organisms highly sensitive to changes                     area burned (Vose and others 2018).                      Efforts to lower hazards focus on
in temperature, which influence their                     Suppression costs and risks to homes                     reducing surface fuels, increasing
survival and population growth (Bentz                     and other infrastructure are also                        the height to live crowns, decreasing
and others 2010). Drought stress                          increasing (Flannigan and others 2006).                  crown bulk density, and retaining large
adversely affects the ability of conifers                                                                          trees of fire-resistant species such as

Fire Management Today                                                           15                                          JANUARY 2021 • VOL. 79 • NO. 1
Bark Beetle and Fire Interactions in Western Coniferous Forests: Research Findings
Fettig and McKelvey (2014) monitored
A common management concern is that bark beetles                                              the effects of fuel reduction treatments
                                                                                              on levels of tree mortality at Blacks
might colonize and kill trees injured by prescribed fire.                                     Mountain Experimental Forest in
                                                                                              California over a 10-year period. Twelve
                                                                                              experimental plots (ranging from
                                                                                              190 to 356 acres (76–142 ha)) were
 ponderosa pine (Agee and Skinner             and others (2006) showed that chipping          established to create two distinct forest
 2005). When applied under prescription,      submerchantable and unmerchantable              structural types: midseral stage (with
 planned ignitions and their mechanical       ponderosa pines and depositing the              low structural diversity) and late-seral
 surrogates (such as thinning from below)     chips back into treated stands increases        stage (with high structural diversity).
 are generally effective in meeting fuel      the risk of infestation by several species      Following harvesting, half of each plot
 reduction goals (McIver and others           of bark beetles in the Southwestern             was treated with prescribed fire.
 2013; Stephens and others 2012). For         United States. The effect was due
 example, the effectiveness of prescribed     to large amounts of monoterpenes                A total of 16,473 trees (9 percent of all
 fire for treating surface and ladder         being released during chipping, which           trees) died. Mortality was concentrated:
 fuels to reduce the incidence of passive     enhanced attraction to bark beetles.
                                                                                              z On plots with high structural diversity
 crown fire (that is, the torching of small   Impacts were greater from chipping
                                                                                                (64 percent);
 groups of trees) is well supported by        in spring (April–May) than in late
 modeling of predicted fire behaviors         summer (August–September) because               z On burned-split plots (61 percent);
 (Stephens and others 2009) and by            spring is the time of peak flight activity      z Within the two smallest diameter
 empirical research (Ritchie and others       for several species of bark beetles in            classes (87 percent); and
 2007). Furthermore, results from the         the Southwestern United States as
                                                                                              z During the second sample period (3 to
 National Fire and Fire Surrogate Study,      they search for new hosts. If possible,
                                                                                                5 years after prescribed burns).
 the largest study of its kind (www.          chipping should be conducted in fall to
 frames.gov/ffs/about), indicate that the     minimize tree losses to bark beetles if         Most mortality was caused by bark
 incidence of active crown fire is best       the chips will remain onsite.                   beetles (65 percent), notably fir
 reduced by combining prescribed fire                                                         engraver (Scolytus ventralis) in white
 with mechanical fuel treatments (McIver      PRESCRIBED FIRE                                 fir (Abies concolor) and mountain pine
 and others 2013).                            Following fire, tree mortality can be           beetle, western pine beetle, and pine
                                              immediate due to consumption of                 engraver (Ips pini) in ponderosa pine.
 The type of fuel reduction treatments                                                        The authors concluded that this level
                                              living tissue or heating of critical plant
 and their manner of implementation                                                           of tree mortality did not interfere
                                              tissues; or it can be delayed, occurring
 have different effects on the fuel matrix,                                                   with management objectives aimed at
                                              over the course of a few years as a
 which can influence the susceptibility of                                                    increasing overall forest resilience.
                                              result of fire injuries to the crown, bole,
 forests to bark beetles in different ways
                                              or roots (Hood and others 2018a).               Similarly, Douglas-fir beetle, pine
 (Fettig and others 2007). For example,
                                              Levels of delayed tree mortality caused         engraver, and western pine beetle caused
 prescribed fire can affect the health
                                              by bark beetles depend on numerous              some tree mortality following prescribed
 and vigor of residual trees; the size,
                                              factors, including tree species; tree size;     fires in western Montana. Mortality
 distribution, and abundance of preferred
                                              tree phenology; degree of fire-caused           occurred shortly after prescribed fires,
 bark beetle hosts; and the physical
                                              injuries; initial and postfire levels of tree   and unburned plots were unaffected.
 environment within forests. Associated
                                              vigor; the postfire environment; and            However, following a regional mountain
 reductions in tree density can alter
                                              the scale, severity, and composition of         pine beetle outbreak that started about 5
 microclimates, affecting beetle fecundity
                                              bark beetle populations and other tree          years after treatments were completed,
 (the ability to produce offspring) and
                                              mortality agents in the area.                   50 percent of ponderosa pines in control
 fitness as well as the phenology (timing
 of life cycle events) and voltinism          A common management concern is that             (untreated) plots and 39 percent in
 (number of generations per unit of time)     bark beetles might colonize and kill trees      prescribe-burned plots were colonized
 of bark beetles and their predators,         that were injured by prescribed fire and
 parasites, and competitors. Tree density     otherwise would have survived. These
 reductions can also disrupt pheromone        trees may then serve as a source of                It is reasonable to
 plumes that attract bark beetles to a tree   beetles and attractive semiochemicals as
 during initial colonization.                 host volatiles are released by the boring          assume that bark
                                              activity of bark beetles. In addition to           beetles and wildfires will
 Volatiles (volatile organic compounds)
                                              host volatiles, the pheromones produced            increasingly interact to
 released from trees are known to
                                              by bark beetles might attract other                shape western forests.
 influence the behavior of many bark
                                              beetles and result in additional levels of
 beetles (Seybold and others 2006). Fettig
                                              tree mortality over time.

 Fire Management Today                                           16                                  JANUARY 2021 • VOL. 79 • NO. 1
Bark Beetle and Fire Interactions in Western Coniferous Forests: Research Findings
and killed. Almost no trees were killed        that unburned areas do not benefit from      2006). Six and others (2002) showed that
by mountain pine beetle in thinned             the positive effects of prescribed fire      pine engravers are unable to colonize
plots and thinned-and-prescribe-burned         (such as increased growing space due to      and reproduce in chips. Moreover,
plots (Hood and others 2016). Thinning         reduced tree density), which affect tree     slash can be managed to minimize
treatments, with or without prescribed         vigor and susceptibility to colonization     colonization of residual trees by bark
fire, dramatically increased tree growth       by bark beetles. Notable infestations in     beetles (DeGomez and others 2015).
rates and production of resin ducts            adjacent unburned areas are uncommon
(a measure of conifer defense against          but can occur and should be watched          EFFECTS OF WILDFIRES ON
bark beetles) relative to the control and      for in case additional management is         BARK BEETLES
prescribed fire treatments.                    warranted to limit tree losses (Fettig and   Factors that influence tree mortality
                                               Hilszczański 2015).                          caused by bark beetles are the same
In some cases, concerns about
maintaining large-diameter trees                                                            after wildfires as after prescribed fires.
                                               MECHANICAL FUEL                              Our distinction is based not on ignition
following prescribed fire have been
justified. For example, Fettig and
                                               TREATMENTS                                   type but largely on differences in fire
McKelvey (2014) reported that most             Factors such as stand density, host          intensity and fire severity: most wildfires
tree mortality (78 percent) in the largest     density, and average tree diameter are       are higher in intensity and severity than
diameter class occurred during the first       strong predictors of the severity of bark    prescribed fires (though not always).
5 years after prescribed fire and that         beetle infestations in the Western United    Low-severity wildfires can induce tree
66 percent was caused by bark beetles.         States. High levels of beetle-caused         defenses against bark beetles (Hood and
Tree protection treatments (such as            tree mortality (for example, greater         others 2015). Resin-duct-related defenses
insecticides and semiochemicals)               than 20 percent) should be expected          take about 1 year after wildfire to form;
can be selectively used to protect             following fuel reduction treatments          during this time, fire-injured trees can be
individual trees from colonization by
bark beetles (Fettig and Hilszczański
2015). Furthermore, methods such as               Firefighters should anticipate the potential for unusual
raking litter and duff from the bases of          fire behavior in beetle-affected forests and the unique
large-diameter trees have been shown              suppression challenges that can result.
to reduce prescribed fire severity and
levels of tree mortality (Fowler and
others 2010; Hood 2010). Additional
research is needed to determine under          that retain high residual stand densities,   more susceptible to colonization by bark
what conditions large-diameter trees are       regardless of treatment effects. Although    beetles, which might help explain some
most susceptible to delayed mortality          thinning has long been advocated as a        of the near-term increases in levels of
following prescribed fire and when tree        measure to reduce beetle-caused tree         beetle-caused tree mortality after some
protection treatments are warranted.           mortality (Fettig and others 2007),          wildfires and prescribed fires (Hood and
                                               thinning prescriptions for fuel reduction    others 2015, 2016). The level of tree
The limited number of studies on the           differ from prescriptions for reducing       injury influences bark beetle attraction,
effects of season of burn (spring versus       susceptibility to bark beetles. In the       with moderately injured trees being
fall) on levels of tree mortality caused by    latter, crown or selection thinning          most susceptible to colonization by bark
bark beetles show mixed results. Some          (that is, removal of larger trees in the     beetles (see, for example, Hood and
studies show increases in certain bark         dominant and codominant crown                Bentz 2007; Lerch and others 2016; and
beetle species following fall treatments       classes) is typically required to achieve    Powell and others 2012).
(that is, when fuels are drier and burns       target threshold densities and residual
are more intense); see, for example,           tree spacing as well as significant          High-severity wildfires generally reduce
Fettig and others (2010). Other studies        reductions in the abundance of preferred     susceptibility to bark beetles by killing
show stronger effects following early-         hosts. Nevertheless, thinning from           large numbers of host trees. For example,
season burns (that is, when bark beetles       below (for fuels reduction) does release     research in subalpine forests in Colorado
are more active); see, for example,            growing space, reducing a stand’s            shows that spruce beetle outbreaks are
Schwilk and others (2006). More                susceptibility to bark beetles.              reduced for decades after high-severity
research is needed to fully define these                                                    wildfires (Bebi and others 2003), the
relationships in different forest types.       A common concern following                   dominant fire regime in these forests. As
                                               mechanical fuel treatments is that bark      with prescribed fires, bark beetles routinely
Although most of the tree mortality            beetles could breed in logging residues      cause additional levels of tree mortality
caused by bark beetles following               (chips and/or slash) and emerge to           after wildfires, but infestations in adjacent
prescribed fire occurs during the first few    colonize residual trees. However, most       unburned areas are uncommon (Davis
years, this pattern might differ in adjacent   studies indicate that this is uncommon       and others 2012; Lerch and others 2016;
untreated areas. The reason, in part, is       (see, for example, Fettig and others         Powell and others 2012).

Fire Management Today                                            17                                JANUARY 2021 • VOL. 79 • NO. 1
EFFECTS OF BARK BEETLE
 OUTBREAKS ON FUELS
 AND FIRE BEHAVIOR AND
 SEVERITY
 Although fuel reduction treatments
 and wildfires can affect bark beetles,
 the reverse is also true: bark beetles
 can alter wildfire behavior by changing
 fuel conditions. Of the bark beetle–
 host systems to consider, the effects
 of mountain pine beetle in lodgepole
 pine (Pinus contorta) have been most
 intensively studied (table 1), and for
 good reason: mountain pine beetle alone
 is responsible for almost half of the
 total area affected by bark beetles in the
 Western United States. All other bark
 beetle–host systems have received less
 attention, some little or none (table 1).
                                              Figure 2—Bark beetles can cause dramatic changes to forest canopies in coniferous forests.
 Fire behavior in beetle-affected forests     For example, during and after mountain pine beetle (Dendroctonus ponderosae) outbreaks,
 largely depends on the severity of           lodgepole pine (Pinus contorta) canopies transition from the green-infested stage (A) to the
 the outbreak (the proportion of trees        red stage (B) in about 1 to 3 years and then to the gray stage (C), which lasts from about 3 to
 colonized and killed) and the amount         25 years following the outbreak. The likelihood of crown and spot fires is greater during the
 of time since the outbreak occurred.         red stage and possibly reduced during the gray stage. Bark beetle activity frequently results in
 Jenkins and others (2008, 2014) use the      a mosaic of green, red, and gray trees (D) as trees are attacked and killed over several years
 term “bark beetle rotation” to describe      (usually 2 to 6 years for mountain pine beetle), which can complicate fire behavior prediction in
 the period from the start of a bark beetle   these forests. Photos: J. Runyon, USDA Forest Service.
 outbreak to the next outbreak within
 susceptible forests. In the “endemic         trees turn red (fig. 2B). The final stage is       energy required for fuels to ignite and
 phase,” beetle-caused tree mortality         the “gray” stage, when needles fall off            burn. Trees colonized and killed by
 is limited (for example, to less than 2      the trees (fig. 2C). The timing of needle          bark beetles rapidly dry out and lose
 trees per acre per year) and generally       fade and color change varies considerably          most of their water content by the first
 isolated to stressed hosts. In the           by tree species, bark beetle species, and          summer following attack as needles
 “epidemic phase,” beetles colonize and       geographic location. For example, in               transition from green to red (fig. 2A,
 kill large numbers of susceptible hosts.     the mountain pine beetle–lodgepole                 2B). For example, the twigs and needles
 In the “post-epidemic phase,” beetle         pine system, the green-infested stage              of lodgepole pine killed by mountain
 populations subside and most beetle-         lasts for about 1 year; the yellow and red         pine beetle lose 80 to 90 percent of
 killed trees fall to the forest floor. The   stages last for about 1 to 3 years; and the        their water content within 1 year of
 endemic and post-epidemic phases can         gray stage lasts for about 3 to 25 years           attack (Jolly and others 2012a; Page
 last for decades to centuries, whereas the   (Klutsch and others 2009). It is important         and others 2012). The loss of moisture
 epidemic phase usually lasts from 2 to       to note that, during an outbreak, trees            increases flammability by shortening
 10 years.                                    are attacked and killed over several years,        time to ignition, lowering temperature
 Recently attacked trees are referred to      and cumulative levels of tree mortality            at ignition, and raising heat yields when
 as “green-infested” (fig. 2A). As needles    vary considerably even within the same             burned (Jolly and others 2012a; Page
 fade, trees enter the “yellow” stage. In     bark beetle–host system. Therefore, a              and others 2012). Reduction in moisture
 the “red” stage, needles on beetle-killed    forest often contains trees and stands             content explains nearly 80 percent of the
                                              in multiple stages and phases of a bark            increase in needle flammability (Page
                                              beetle rotation at the same time (fig. 2D).        and others 2012). Similar reductions in
                                                                                                 foliage moisture content and increases
Not surprisingly, beetle-                     CROWN AND CANOPY FUELS                             in flammability have been documented
induced changes to foliar                     Not surprisingly, beetle-induced changes           in Engelmann spruce (Picea engelmannii)
                                              to foliar moisture have the greatest               killed by spruce beetle (Page and others
moisture have the greatest                    effects on flammability. This is because           2014) and Douglas-fir killed by Douglas-
effects on flammability.                      water is a heat sink and moisture in               fir beetle (Giunta 2016).
                                              plant tissues increases the amount of

 Fire Management Today                                            18                                      JANUARY 2021 • VOL. 79 • NO. 1
the ground (Hoffman and others 2015).
                                                                                                         Increased light and moisture availability
                                                                                                         resulting from mortality of dominant
                                                                                                         and codominant trees release smaller
                                                                                                         surviving trees that were unsuitable
                                                                                                         hosts for bark beetles, increasing ladder
                                                                                                         fuels during the post-epidemic phase.

                                                                                                         LITTER, FINE FUELS, AND
                                                                                                         COARSE WOODY FUELS
                                                                                                         Although dying and dead foliage is more
                                                                                                         flammable than live foliage, it remains in
                                                                                                         the canopy for only a short time (usually
                                                                                                         3 years or less). The accumulation rate
                                                                                                         of canopy materials (foliage and twigs)
                                                                                                         on the forest floor can be of greater
                                                                                                         importance, influencing surface fuel
                                                                                                         loadings and associated fire hazards.
                                                                                                         Stalling and others (2017) evaluated
                                                                                                         the effects of mountain pine beetle
                                                                                                         and Douglas-fir beetle outbreaks on
                                                                                                         fuel conditions in Montana and Idaho.
                                                                                                         Foliage deposition occurred mostly
                                                                                                         during the first 2 years of the epidemic
Figure 3—Coarse woody fuel accumulations following a bark beetle outbreak in the Sierra                  phase. Unlike foliage, fine woody
Nevada in California. Based on a survey of 180 plots across 4 national forests, 31 percent of the        and nonwoody material tended to
dead trees fell to the forest floor within 4 years of being killed, which is much faster than reported   come down irregularly, deposited at
elsewhere in the literature for other locations and bark beetle–host systems (Fettig and others 2019).   essentially the same rate from year to
Stephens and others (2018) concluded that a greater potential for “mass fires” exists for this region,   year. Tree fall and accumulations of
driven by the amount, size, and continuity of dry combustible woody fuels, which could produce           coarse woody surface fuels were limited
large, severe, and uncontrollable wildfires. Mass fires or “firestorms” (Finney and McAllister 2011)     over the 10-year period of study. These
can occur when large areas burn simultaneously for long periods at high intensities, generating their    findings suggest that fire hazards in these
own weather conditions. The science on mass fires (such as Countryman 1965) is limited and the           forests were influenced less during the
risks are poorly understood. Photo: L. Mortenson, USDA Forest Service.                                   late epidemic and early post-epidemic
                                                                                                         phases than previously thought. Highly
Bark beetles alter foliage flammability in           2011; Courty and others 2012) but has               flammable dead foliage does not stay in
other ways as well. The proportions of               not yet been studied in the Western                 the canopy for long, and accumulations
fat, fiber, lignin and cellulose, starches,          United States. Firefighters have reported           of canopy materials on the forest floor
and sugars change in needles after                   observations that support the existence             do not exceed the annual rate of fuel
beetle colonization, and each factor can             of terpene clouds in western forests                decomposition (Stalling and others 2017).
affect flammability (Jolly and others                during the epidemic phase; additional
2012a; Page and others 2012, 2014).                  research is warranted on terpene                    Ultimately, all of the woody fuel from
Conifers also contain large amounts                  clouds and their potential effects on fire          beetle-killed trees is transferred to the
of flammable terpenes. Beetle attack                 behavior in the Western United States.              forest floor during the post-epidemic
generally increases the emission of                                                                      phase. This period varies considerably
terpenes and terpene concentrations                  Outbreaks also alter canopy fuel                    by site and tree species, among other
within needles (Giunta and others 2016;              arrangement (Hicke and others 2012;                 factors, with half-lives (the time required
Page and others 2012, 2014), which                   Jenkins and others 2008). As trees move             for half of the dead trees to fall to
can shorten time to ignition, lower                  from the red to the gray stage (fig. 2B,            the forest floor) ranging from years
temperature at ignition, and increase                2C), canopy fuels decrease. As dead                 (such as for ponderosa pines killed by
the maximum rate of mass loss (an                    trees deteriorate and fall to the forest            western pine beetle in the central and
indication of burning rate) (Page and                floor (fig. 3), canopy bulk density and             southern Sierra Nevada (fig. 3)) to
others 2012, 2014). Moreover, the                    canopy cover decline over time. The                 decades (such as for lodgepole pines
emission of terpene “clouds” from                    decrease in canopy fuel continuity also             killed by mountain pine beetle in
plants has been linked to eruptive fire              reduces the sheltering effect of the                northeast Oregon (Harvey 1986)). Over
behavior in Europe (Barboni and others               forest, causing higher wind speeds near             time, medium and coarse woody fuels

Fire Management Today                                                     19                                    JANUARY 2021 • VOL. 79 • NO. 1
gradually accumulate, increasing fuel                 Physics-based models suggest that bark              Using a physics-based model, Sieg and
bed depth. For example, Jenkins and                   beetle outbreaks increase wildfire rates of         others (2017) found higher fire severity
others (2014) reported that medium                    spread, with spread rates peaking during            (that is, tree mortality) in ponderosa-
and coarse woody fuels increased by a                 the red stage. Rates of spread remain               pine-dominated forests during the red
factor of about 2.5 to 8 in forests of the            higher than in the endemic phase even               stage than in the endemic phase but
Intermountain West following mountain                 though canopy fuels decrease (Hoffman               unchanged or even lower fire severity
pine beetle outbreaks 20 years earlier.               and others 2015). In an experiment on               during the gray stage. The observed
                                                      needle flammability, red needles from               increases in fire severity attributed to the
FIRE BEHAVIOR                                         beetle-attacked trees ignited faster than           bark beetle outbreak declined under high
AND SEVERITY                                          green needles, which could lead to                  wind conditions (Sieg and others 2017).
Modeling fire behavior in beetle-affected             increased crown fire potential (Jolly and
                                                      others 2012b). However, it is unknown               Fire severity, measured as change
forests is challenging, largely because fire                                                              in vegetation, decreased over time
behavior model assumptions are violated               whether the increase in flammability
                                                      scales up to entire canopies and results in         following mountain pine beetle
(Hood and others 2018b; Jenkins and                                                                       outbreaks in Washington and Oregon
others 2012, 2014; Page and others 2014).             higher intensity crown fires (Alexander
                                                      and Cruz 2013b). After a crown fire                 (Meigs and others 2016). By contrast,
The limited ability to use empirical data                                                                 Prichard and Kennedy (2014) reported
to evaluate model predictions (Alexander              begins, fire behavior in lodgepole pine
                                                      forests under dry, windy conditions is              higher fire severity during the red
and Cruz 2013a) has fed controversy over                                                                  stage following mountain pine beetle
whether epidemic and/or post-epidemic                 likely to be similar, regardless of bark
                                                      beetle activity (Schoennagel and others             outbreaks in Washington.
phases have more fire behavior hazards
than the endemic phase (Jolly and others              2012). Under most circumstances, severe             Harvey and others (2014) reported
2012a; Simard and others 2011, 2012).                 fire weather conditions trump beetle-               similar levels of fire severity in red-
                                                      induced changes in fuel conditions.                 stage and gray-stage lodgepole pine
                                                                                                          forests following mountain pine beetle
                                                                                                          outbreaks in the Northern Rockies.
                                                                                                          They evaluated several metrics of fire
                                                                                                          severity and found that only extreme
                                                                                                          fire weather conditions increased fire
                                                                                                          severity in terms of deep charring (that
                                                                                                          is, charring into the wood along tree
                                                                                                          boles and into crowns, with less than 5
                                                                                                          percent of branches remaining) (Harvey
                                                                                                          and others 2014).
                                                                                                          After forests enter the gray stage,
                                                                                                          evidence suggests that fire severity—
                                                                                                          and, presumably, the potential for
                                                                                                          heightened fire behavior—diminish for
                                                                                                          some time (Hicke and others 2012) (fig.
                                                                                                          4). However, more research is needed to
                                                                                                          fully understand the potential for crown
                                                                                                          fires in the gray stage.
                                                                                                          Although bark beetle outbreaks can
                                                                                                          influence fire behavior and severity, they
                                                                                                          have little effect on the extent of the area
                                                                                                          burned (see, for example, Hart and others
                                                                                                          2015) or the likelihood or frequency of
                                                                                                          wildfire occurrence (Bebi and others
                                                                                                          2003; Meigs and others 2015).

Figure 4—Gray-stage coniferous forest following a bark beetle outbreak. Fire behavior in                  The effects on fire behavior and severity
beetle-affected forests largely depends on the proportion of trees killed and the amount of time          have important implications for fire
since the bark beetle outbreak (that is, the “bark beetle rotation”). In the gray stage, fire severity—   management and firefighter safety. For
and, presumably, the potential for heightened fire behavior—is reduced. However, high-severity            example, the likelihood of torching,
crown fires have been reported in some gray-stage forests (Agne and others 2016), especially after        crowning, and spotting can be greater in
significant accumulations of coarse woody fuels. The science on fire behavior in gray-stage forests is    forests containing an abundance of red
limited. Photo: S. Hood, USDA Forest Service.

Fire Management Today                                                      20                                    JANUARY 2021 • VOL. 79 • NO. 1
crowns (Schoennagel and others 2012;         fire interactions in western coniferous          small trees or big trees are killed and
 Jenkins and others 2014). Moreover,          forests in the last 15 years. Much of            whether some trees or nearly all trees
 firefighters have observed “surprising”      this information has been synthesized            are killed);
 fire behavior in some forests during the     in notable publications (such as Hicke         z The dominant fire regime of the forest
 epidemic and post-epidemic phases            and others 2012; Jenkins and others              type;
 (Moriarty and others 2019; Page and          2008, 2012, 2014; Kane and others
 others 2013). Rapid shifts from surface      2017; and Stephens and others 2018).           z The spatial scale; and
 fires to crown fires pose safety risks       We encourage the reader interested in          z Limitations in the prediction of fire
 to firefighters caused by increases in       delving deeper into this topic to consult        behavior in beetle-affected forests.
 fireline intensities and spotting, which     these publications.                            Furthermore, most studies are
                                                                                             retrospective (rather than controlled
                                                                                             experiments) and inherently have a lot
                                                                                             of variability.
Since 2000, about 25.5 million acres in the Western United
States have been affected by mountain pine beetle.                                           Given the implications for firefighter
                                                                                             safety and fire suppression activities,
                                                                                             work is needed to more accurately
                                                                                             quantify relationships between bark
                                                                                             beetle outbreaks and fire behavior and
 might necessitate larger safety zones        The scientific community now has               severity, especially in understudied
 (Butler and Cohen 1998). Snags are an        a pretty solid understanding of the            forest types (table 1). Creating and
 important safety concern, particularly       effects of fuel reduction treatments           maintaining forest structures that
 during the post-epidemic phase (fig.         on bark beetles, which tend to lead            are more resilient to bark beetles and
 3). Accordingly, firefighters should         to fairly consistent responses among           wildfires will go a long way toward
 anticipate the potential for unusual fire    different forest types in the Western          addressing concerns regarding increases
 behavior (even during the green-infested     United States. Nevertheless, knowledge         in fire behavior and severity following
 stage) in beetle-affected forests and the    gaps exist and need to be addressed.           bark beetle outbreaks.
 unique suppression challenges that can       Many initial fears concerning long-
 result (such as increased difficulties in    term increases in levels of delayed            ACKNOWLEDGMENTS
 fireline construction and establishment      tree mortality caused by bark beetles          The genesis of this article was the
 of access, egress, and escape routes).       were unfounded. Accordingly, one               symposium “Bark Beetle and Fire
                                              might view the associated increases in         Interactions in Western North America:
 LOOKING FORWARD                              tree mortality as “short-term losses”          The Current State of Knowledge
                                              suffered for “long-term gains” (Fettig         and Implications for Forest and Fire
 A key finding of the recently published
                                              and McKelvey 2014). This is especially         Managers,” chaired by Christopher
 U.S. Fourth National Climate
                                              true when considering that rates of            J. Fettig and Justin B. Runyon at the
 Assessment is that it is “very likely
                                              tree mortality caused by bark beetles          7th International Fire Congress held
 that more frequent extreme weather
                                              are generally low (less than 5 percent)        in Orlando, FL, in November 2017.
 events will increase the frequency
                                              and concentrated in small-diameter             We acknowledge the support of—and
 and magnitude of severe ecological
                                              trees (that is, ladder fuels) and in fire-     the thoughtful debate and dialogue
 disturbances, driving rapid (months to
                                              intolerant tree species (such as white fir).   conducted by—numerous colleagues at
 years) and often persistent changes in
 forest structure and function across large   Although the scientific literature shows       that symposium and elsewhere, which
 landscapes” (Vose and others 2018). As       mixed results, most suggests that bark         helped shaped the content of this article
 such, it is reasonable to assume that bark   beetle outbreaks can significantly             and furthered our understanding of bark
 beetles and wildfires will increasingly      change fuel profiles and fire behavior         beetle and fire interactions. We thank Dr.
 interact to shape western forests.           and severity during the epidemic and           Daniel Dey (USDA Forest Service) for
                                              post-epidemic phases. Nevertheless,            the invitation to contribute to this special
 Recognizing this, the fire science and                                                      issue and Dr. Dey and Hutch Brown
                                              discussions of the effects of bark beetle
 forest health communities have largely                                                      (USDA Forest Service) for their reviews.
                                              outbreaks on fuels and fire behavior get
 bridged cultural and communication                                                          Much of the research cited herein was
                                              unnecessarily complex and occasionally
 divides (Jenkins and others 2009) and                                                       funded, in part, by the National Fire and
                                              contentious if they ignore:
 have strengthened relationships with                                                        Fire Surrogate Study, the Forest Health
 managers in hopes of delivering science      z The amount of time that has occurred         Monitoring-Evaluation Monitoring
 of utmost relevance to managers’               since the outbreak (that is, the bark        Program, and the USDI-USDA Joint
 concerns (Kocher and others 2012).             beetle rotation);                            Fire Sciences Program.
 A tremendous amount of knowledge
                                              z The type and severity of the outbreak
 has been developed on bark beetle and
                                                (including such factors as whether

 Fire Management Today                                           21                                 JANUARY 2021 • VOL. 79 • NO. 1
LITERATURE CITED                                            In: Vega, F.E.; Hofstetter, R.W. eds. Bark        Harvey, R.D. 1986. Deterioration of mountain
                                                            beetles: biology and ecology of native and            pine beetle-killed lodgepole pine in northeast
Agee, J.K.; Skinner, C.N. 2005. Basic principles
                                                            invasive species. London: Springer: 555–584.          Oregon. Pap. R6–86–13. Portland, OR:
    of forest fuel reduction treatments. Forest
                                                        Fettig, C.J.; McMillin, J.D.; Anhold, J.A. [and           USDA Forest Service. 10 p.
    Ecology and Management. 211: 83–96.
                                                            others]. 2006. The effects of mechanical          Hicke, J.A.; Johnson, M.C.; Hayes, J.L.; Preisler,
Agne, M.C.; Woolley, T.; Fitzgerald, S. 2016.
                                                            fuel reduction treatments on the activity             H.K. 2012. Effects of bark beetle-caused tree
    Fire severity and cumulative disturbance
                                                            of bark beetles (Coleoptera: Scolytidae)              mortality on wildfire. Forest Ecology and
    effects in the post-mountain pine beetle
                                                            infesting ponderosa pine. Forest Ecology              Management. 271: 81–90.
    lodgepole pine forests of the Pole Creek
                                                            and Management. 230: 55–68.                       Hicke, J.A.; Meddens, A.J.H.; Kolden, C.A.
    Fire. Forest Ecology and Management. 366:
    73–86.                                              Fettig, C.J.; Klepzig, K.D.; Billings, R.F. [and          2016. Recent tree mortality in the Western
                                                            others]. 2007. The effectiveness of vegetation        United States from bark beetles and forest
Alexander, M.E.; Cruz, M.G. 2013a.
                                                            management practices for prevention                   fires. Forest Science. 62: 141–153.
    Limitations on the accuracy of model
                                                            and control of bark beetle infestations           Hoffman, C.M.; Linn, R.; Parsons, R. [and
    predictions of wildland fire behaviour:
                                                            in coniferous forests of the western and              others]. 2015. Modeling spatial and
    a state-of-the-knowledge overview. The
                                                            southern United States. Forest Ecology and            temporal dynamics of wind flow and
    Forestry Chronicle. 89: 372–383.
                                                            Management. 238: 24–53.                               potential fire behavior following a mountain
Alexander, M.E.; Cruz, M.G. 2013b. Assessing
                                                        Fettig, C.J.; McKelvey, S.R.; Cluck, D.L. [and            pine beetle outbreak in a lodgepole pine
    the effect of foliar moisture on the spread
                                                            others]. 2010. Effects of prescribed fire and         forest. Agricultural and Forest Meteorology.
    rate of crown fires. International Journal of
                                                            season of burn on direct and indirect levels          204: 79–93.
    Wildland Fire. 22: 415–427.
                                                            of tree mortality in ponderosa and Jeffrey        Hood, S.M. 2010. Mitigating old tree mortality
Barboni, T.; Cannac, M.; Leoni, E.;                         pine forests in California, USA. Forest               in long-unburned, fire-dependent forests:
    Chiaramonti, N. 2011. Emission of biogenic              Ecology and Management. 260: 207–218.                 a synthesis. Gen. Tech. Rep. RMRS–
    volatile organic compounds involved in
                                                        Fettig, C.J.; Mortenson, L.A.; Bulaon, B.M.;              GTR–238. Fort Collins, CO: USDA Forest
    eruptive fire: implications for the safety
                                                            Foulk, P.B. 2019. Tree mortality following            Service, Rocky Mountain Research Station.
    of firefighters. International Journal of
                                                            drought in the central and southern Sierra            71 p.
    Wildland Fire. 20: 152–161.
                                                            Nevada, California, U.S. Forest Ecology and       Hood, S.M.; Bentz, B. 2007. Predicting post-
Bebi, P.; Kulakowski, D.; Veblen, T.T. 2003.                Management. 432: 164–178.                             fire Douglas-fir beetle attacks and tree
    Interactions between fire and spruce beetles
                                                        Finney, M.A.; McAllister, S.S. 2011. A review             mortality in the Northern Rocky Mountains.
    in a subalpine Rocky Mountain forest
                                                            of fire interactions and mass fires. Journal          Canadian Journal of Forest Research. 37:
    landscape. Ecology. 84: 362–371.
                                                            of Combustion. 2011: Article 548328. DOI:             1058–1069.
Bentz, B.J.; Régnière, J.; Fettig, C.J. [and others].       10.1155/2011/548328.                              Hood, S.; Sala, A.; Heyerdahl, E.K.; Boutin, M.
    2010. Climate change and bark beetles of the
                                                        Flannigan, M.D.; Amiro, B.D.; Logan, K.A.                 2015. Low-severity fire increases tree defense
    Western United States and Canada: direct
                                                            [and others]. 2006. Forest fires and climate          against bark beetle attacks. Ecology. 96:
    and indirect effects. Bioscience. 60: 602–613.
                                                            change in the 21st century. Mitigation and            1846–1855.
Butler, B.W.; Cohen, J.D. 1998. Firefighter safety          Adaptation Strategies for Global Change.          Hood, S.; Baker, S.; Sala, A. 2016. Fortifying
    zones: a theoretical model based on radiative           11: 847–859.                                          the forest: thinning and burning increase
    heating. International Journal of Wildland
                                                        Fowler, J.F.; Sieg, C.H.; Wadleigh, L.L. 2010.            resistance to a bark beetle outbreak and
    Fire. 8: 73–77.
                                                            Effectiveness of litter removal to prevent            promote forest resilience. Ecological
Countryman, C.M. 1965. Mass fire                            cambial kill-caused mortality in northern             Applications. 26: 1984–2000.
    characteristics in large-scale tests. Fire              Arizona ponderosa pine. Forest Science. 56:       Hood, S.; Varner, M.; van Mantgem, P.;
    Technology. 1: 303–317.                                 166–171.                                              Cansler, C.A. 2018a. Fire and tree death:
Courty, L.; Chetehouna, K.; Halter, F. [and             Giunta, A. 2016. Douglas-fir beetle mediated              understanding and improving modeling of
    others]. 2012. Flame speeds of α-pinene/                changes to fuel complexes, foliar moisture            fire-induced tree mortality. Environmental
    air and limonene/air mixtures involved in               content, and terpenes in interior Douglas-            Research Letters. 13: 113004.
    accelerating forest fires. Combustion Science           fir forests of the central Rocky Mountains.       Hood, S.; Keane, R.E.; Smith, H.Y. [and others].
    and Technology. 184: 1397–1411.                         Master’s Thesis. Logan, UT: Utah State                2018b. Conventional fire behavior modeling
Davis, R.S.; Hood, S.; Bentz, B.J. 2012. Fire-              University. 161 p.                                    systems are inadequate for predicting fire
    injured ponderosa pine provide a pulsed             Giunta, A.D.; Runyon, J.B.; Jenkins, M.J.; Teich,         behavior in bark beetle-impacted forests. In:
    resource for bark beetles. Canadian Journal             M. 2016. Volatile and within-needle terpene           Potter, K.; Conkling, B., eds. Forest health
    of Forest Research. 42: 2022–2036.                      changes to Douglas-fir trees associated               monitoring: national status, trends, and
DeGomez, T.; Fettig, C.J.; McMillin, J.D.;                  with Douglas-fir beetle (Coleoptera:                  analysis 2017. Gen. Tech. Rep. SRS–GTR–
    Anhold, J.A.; Hayes, C. 2015. Managing                  Curculionidae) attack. Environmental                  233. Asheville, NC: USDA Forest Service,
    slash to minimize colonization of residual              Entomology. 45: 920–929.                              Southern Research Station: 167–176.
    leave trees by Ips and other bark beetle            Hart, S.J.; Schoennagel, T.; Veblen, T.T.;            Jenkins, M.J.; Hebertson, E.; Page, W.;
    species following thinning in southwestern              Chapman, T.B. 2015. Area burned in the                Jorgensen, W.P. 2008. Bark beetles, fuels,
    ponderosa pine (revised 10/2014). AZ1449.               Western United States is unaffected by                fires and implications for forest management
    Tucson, AZ: University of Arizona, College              recent mountain pine beetle outbreaks.                in the Intermountain West. Forest Ecology
    of Agriculture and Life Sciences Bulletin.              Proceedings of the National Academy of                and Management. 254: 16–34.
    12 p.                                                   Sciences. 112: 4375–4380.                         Jenkins, M.J.; Fettig, C.J.; Hebertson, E.G. 2009.
Fettig, C.J.; McKelvey, S.R. 2014. Resiliency           Harvey, B.J.; Donato, D.C.; Turner, M.G.                  Bark beetles, fuels and fire: a synthesis of
    of an interior ponderosa pine forest to bark            2014. Recent mountain pine beetle                     our present understanding and implications
    beetle infestations following fuel-reduction            outbreaks, wildfire severity, and postfire tree       for management. In: Rideout-Hanzak,
    and forest-restoration treatments. Forests. 5:          regeneration in the U.S. Northern Rockies.            S.; Oswald, B.P.; Beierle, M., comps. 4th
    153–176.                                                Proceedings of the National Academy of                Annual International Congress on Fire
Fettig, C.J.; Hilszczański, J. 2015. Management             Sciences. 111: 15120–15125.                           Ecology and Management: Fire as a Global
    strategies for bark beetles in conifer forests.

Fire Management Today                                                         22                                      JANUARY 2021 • VOL. 79 • NO. 1
Process. Leavenworth, WA: The Association         Meigs, G.W.; Campbell, J.L.; Zald, H.S.J. [and           Seybold, S.J.; Huber, D.P.W.; Lee, J.C.;
     for Fire Ecology.                                     others]. 2015. Does wildfire likelihood                  Graves, A.D.; Bohlmann, J. 2006. Pine
Jenkins, M.J.; Page, W.G.; Hebertson, E.G.;                increase following insect outbreaks in conifer           monoterpenes and pine bark beetles: a
     Alexander, M.E. 2012. Fuels and fire                  forests? Ecosphere. 6: art118.                           marriage of convenience for defense and
     behavior dynamics in bark beetle-attacked         Meigs, G.W.; Zald, H.S.J.; Campbell, J.L. [and               chemical communication. Phytochemistry
     forests in Western North America and                  others]. 2016. Do insect outbreaks reduce                Reviews. 5: 143–178.
     implications for fire management. Forest              the severity of subsequent forest fires?             Sieg, C.H.; Linn, R.R.; Pimont, F. [and others].
     Ecology and Management. 275: 23–34.                   Environmental Research Letters. 11: 045008.              2017. Fires following bark beetles: factors
Jenkins, M.J.; Runyon, J.B.; Fettig, C.J. [and         Moriarty, K.; Cheng, A.S.; Hoffman, C.M. [and                controlling severity and disturbance
     others]. 2014. Interactions among the                 others]. 2019. Firefighter observations of               interactions in ponderosa pine. Fire Ecology.
     mountain pine beetle, fires, and fuels. Forest        “surprising” fire behavior in mountain pine              13: 1–23.
     Science. 60: 489–501.                                 beetle-attacked lodgepole pine forests. Fire.        Simard, M.; Romme, W.H.; Griffin, J.M.;
Jolly, W.M.; Parsons, R.A.; Hadlow, A.M.                   2: 34. DOI: 10.3390/fire2020034.                         Turner, M.G. 2011. Do mountain pine
     [and others]. 2012a. Relationships between        Morris, J.L.; Cottrell, S.; Fettig, C.J. [and others].       beetle outbreaks change the probability of
     moisture, chemistry, and ignition of Pinus            2018. Bark beetles as agents of change                   active crown fire in lodgepole pine forests?
     contorta needles during the early stages of           in social-ecological systems. Frontiers in               Ecological Monographs. 81: 3–24.
     mountain pine beetle attack. Forest Ecology           Ecology and the Environment. 16(S1):                 Simard, M.; Romme, W.H.; Griffin, J.M.;
     and Management. 269: 52–59.                           S34–S43.                                                 Turner, M.G. 2012. Do mountain pine
Jolly, W.M.; Parsons, R.; Varner, J.M. [and            Page, W.G.; Jenkins, M.J.; Runyon, J.B. 2012.                beetle outbreaks change the probability of
     others]. 2012b. Comment: Do mountain                  Mountain pine beetle attack alters the                   active crown fire in lodgepole pine forests?
     pine beetle outbreaks change the probability          chemistry and flammability of lodgepole                  Reply. Ecology. 93: 946–950.
     of active crown fire in lodgepole pine forests?       pine foliage. Canadian Journal of Forest             Six, D.L.; Vander Meer, M.; DeLuca, T.H.;
     Ecology. 93: 941–946.                                 Research. 42: 1631–1647.                                 Kolb, P. 2002. Pine engraver (Ips pini)
Kane, J.M.; Varner, J.M.; Metz, M.R.; van              Page, W.G.; Alexander, M.E.; Jenkins, M.J.                   colonization of logging residues created
     Mantgem, P.J. 2017. Characterizing                    2013. Wildfire’s resistance to control in                using alternative slash management systems
     interactions between fire and other                   mountain pine beetle-attacked lodgepole                  in western Montana. Western Journal of
     disturbances and their impacts on tree                pine forests. The Forestry Chronicle. 89:                Applied Forestry. 17: 96–100.
     mortality in western U.S. forests. Forest             783–794.                                             Stalling, C.; Keane, R.E.; Retzlaff, M.
     Ecology and Management. 405: 188–199.             Page, W.G.; Jenkins, M.J.; Runyon, J.B.                      2017. Surface fuel changes after severe
Kocher, S.D.; Toman, E.; Trainor, S.F. [and                2014. Spruce beetle-induced changes to                   disturbances in northern Rocky Mountain
     others]. 2012. How can we span the                    Engelmann spruce foliage flammability.                   ecosystems. Forest Ecology and
     boundaries between wildland fire science              Forest Science. 60: 691–702.                             Management. 400: 38–47.
     and management in the United States?              Powell, E.N.; Townsend, P.A.; Raffa, K.F.                Stephens, S.L.; Moghaddas, J.J.; Edminister, C.
     Journal of Forestry. 110: 421–428.                    2012. Wildfire provides refuge from                      [and others]. 2009. Fire treatment effects on
Kolb, T.E.; Fettig, C.J.; Ayres, M.P. [and others].        local extinction but is an unlikely driver               vegetation structure, fuels, and potential fire
     2016. Observed and anticipated impacts                of outbreaks by mountain pine beetle.                    severity in western U.S. forests. Ecological
     of drought on forest insects and diseases             Ecological Monographs. 82: 69–84.                        Applications. 19: 305–320.
     in the United States. Forest Ecology and          Prichard, S.J.; Kennedy, M.C. 2014. Fuel                 Stephens, S.L.; McIver, J.D.; Boerner, R.E.J.
     Management. 380: 321–334.                             treatments and landform modify landscape                 [and others]. 2012. The effects of forest fuel-
Klutsch, J.G.; Negron, J.F.; Costello, S.L.                patterns of burn severity in an extreme fire             reduction treatments in the United States.
     [and others]. 2009. Stand characteristics             event. Ecological Applications. 24: 571–590.             Bioscience. 62: 549–560.
     and downed woody debris accumulations             Ritchie, M.W.; Skinner, C.N.; Hamilton, T.A.             Stephens, S.L.; Collins, B.M.; Fettig, C.J. [and
     associated with a mountain pine beetle                2007. Probability of tree survival after                 others]. 2018. Drought, tree mortality, and
     (Dendroctonus ponderosae Hopkins)                     wildfire in an interior pine forest of northern          wildfire in forests adapted to frequent fire.
     outbreak in Colorado. Forest Ecology and              California: effects of thinning and prescribed           Bioscience. 68: 77–88.
     Management. 258: 641–649.                             fire. Forest Ecology and Management. 247:            Vose, J.M.; Peterson, D.L.; Domke, G.M. [and
Lerch, A.P.; Pfammatter, J.A.; Bentz, B.J.;                200–208.                                                 others]. 2018. Forests. In: Reidmiller, D.R.;
     Raffa, K.F. 2016. Mountain pine beetle            Schoennagel, T.; Veblen T.T.; Negron J.F.;                   Avery, C.W.; Easterling, D.R. [and others],
     dynamics and reproductive success in post-            Smith, J.M. 2012. Effects of mountain pine               eds. Impacts, risks, and adaptation in the
     fire lodgepole and ponderosa pine forests in          beetle on fuels and expected fire behavior in            United States: fourth national climate
     northeastern Utah. Plos One 11: e0164738.             lodgepole pine forests, Colorado, USA. Plos              assessment, volume II. Washington, DC:
McIver, J., Stephens, S.; Agee, J. [and others].           One. 7(1): e30002.                                       U.S. Global Change Research Program:
     2013. Ecological effects of alternative fuel      Schwilk, D.W.; Knapp, E.E.; Ferrenberg, S.M.                 223–258.
     reduction treatments: highlights of the               [and others]. 2006. Tree mortality from fire                                                         ■
     national fire and fire surrogate study (FFS).         and bark beetles following early and late
     International Journal of Wildland Fire. 22:           season prescribed fires in a Sierra Nevada
     63–82.                                                mixed-conifer forest. Forest Ecology and
                                                           Management. 232: 36–45.

Fire Management Today                                                         23                                        JANUARY 2021 • VOL. 79 • NO. 1
You can also read