J. For.

115(5):379 –384
PRACTICE OF FORESTRY https://doi.org/10.5849/jof.16-049

silviculture

Dwarf Mistletoe Control on the Mescalero

Apache Indian Reservation, New Mexico

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Howard M. Hoyt, William Hornsby, Ching-Hsun Huang,
James J. Jacobs, and Robert L. Mathiasen

Dwarf mistletoe is a destructive pathogen of many commercial ponderosa pine (Pinus ponderosa var. ponderosa var. scopulorum Engelmann) for-
scopulorum) stands in the Southwest. From 2005 to 2006, on the Mescalero Apache Indian Reservation, a ests (Hawksworth and Wiens 1996, Conk-
mistletoe control project was implemented across 3,300 acres. Locally developed management guidelines for lin 2000). According to Hawksworth and
treating moderate to severely infected stands recommended salvaging merchantable timber and then slashing Wiens (1996), this dwarf mistletoe is partic-
remaining hosts ⬎2 ft tall. The goal of this treatment was to diminish residual southwestern dwarf mistletoe ularly damaging to ponderosa pine in the
(Arceuthobium vaginatum subsp. cryptopodum) infection while maintaining sufficient, albeit small, natural Sacramento Mountains in southcentral
ponderosa pine regeneration for stocking. In 2008, 12 fixed-radius plots were established within the project area New Mexico, central Arizona, and along the
to monitor the emergence of latent infections. Our observations revealed that 9 years posttreatment, 2.3% of Front Range in Colorado. Shields (1953)
the retained cohort had harbored latent mistletoe infections. The treatment appears to result in satisfactory contended that the most widespread and
control of the parasite in the Sacramento Mountains of southcentral New Mexico. severe infestation of SWDM was on the
Mescalero Apache Indian Reservation (Mes-
Keywords: southwestern dwarf mistletoe, stand improvement, sanitation cut, advanced regeneration,
calero) in southern New Mexico. In addi-
ponderosa pine
tion, the occurrence of SWDM has been
shown to increase surface and total fuel load-
ing in stands and may increase the risk of

D
warf mistletoes (Arceuthobium
spp.) are obligate parasitic flower-
ing plants that depend almost en-
tirely on their host tree for water, minerals,
and carbohydrates. These plants have the
potential to significantly impact height
growth, diameter increment, and life expec-
tancy of a severely infected tree. Dwarf mis-
tletoes are spread within and between
crowns of adjacent trees by hydrostatically
controlled, thermogenesis-triggered, explo-
sive dispersal of their seeds (Hawksworth
and Wiens 1996, Geils et al. 2002, Rolena et
al. 2015). Because of this unique dispersal
mechanism, distribution on both the land-
scape and stand scale is typically patchy with
more or less discrete infection centers sur-
rounded by areas without the disease (Conk-
lin 2000).
Southwestern dwarf mistletoe (SWDM)
(Arceuthobium vaginatum subsp. cryptopo-
dum [Engelmann] Hawksworth and Wiens)
is recognized as the most damaging patho-
gen in southwestern ponderosa pine (Pinus
catastrophic wildfire when the infestation is
severe (Hoffman et al. 2007).
Several aspects of the dwarf mistletoe
life cycle facilitate their control through sil-
vicultural methods including (1) obligate
parasitism—requirement of a living host,
(2) host specificity— generally infect one
species or a group of related species, (3) long
life cycles— generations range from 2–10
years or more including an incubation pe-
riod of 2–10 years before aerial shoots de-

Received August 15, 2016; accepted September 12, 2016; published online November 24, 2016.
Affiliations: Howard M. Hoyt (hhoyt@fs.fed.us), USDA Forest Service, Siuslaw National Forest, Central Coast Ranger District, Waldport, OR. William Hornsby
(william.hornsby@bia.gov), USDI Bureau of Indian Affairs, Mescalero Agency. Ching-Hsun Huang (ching.huang@nau.edu), Northern Arizona University. James J. Jacobs
(jamesjjacobs@fs.fed.us), USDA Forest Service, Forest Health and Protection. Robert L. Mathiasen (robert.mathiasen@nau.edu), Northern Arizona University.
Acknowledgments: We acknowledge the researchers and managers at Mescalero who paved the way for the current study and whose work has been instrumental in the
management of mistletoe at Mescalero, notably the late Frank G. Hawksworth; Dave Conklin, USFS Forest Pathologist (retired); John Andrews, Bureau of Indian
Affairs (BIA) Forester (retired); Bernie Ryan, BIA Forester (retired); Robert Campbell, BIA Forester; Dave Koch, BIA Forester; Thora Padilla, Mescalero Apache
Tribe (MAT) Forester; and Mark Paul, BIA Forester. We also acknowledge the BIA, Mescalero Apache Tribe and USDA Forest Service staff who have worked on
this project, particularly Gerald Blake, Jr., Roderick Chimal, Rylee Chino, George Gallardo, Clay Garrison, Andy Graves, Serra Hoagland, Wilmer LaPaz, Gerald
Magoosh, Phil Smith, Taylor Smith and Crystal Tischler. Funding for this project was obtained from the USDA Forest Service, Forest Health and Protection,
Southwestern Region and the National Fire Plan, Hazardous Fuel Reduction Program.

Journal of Forestry • September 2017 379

velop, (4) limited seed dispersal—maximum
horizontal spread distances of 33– 49 ft, and
(5) slow intensification within crowns—fast
growing trees may outpace internal spread
(Baranyay et al. 1971, Muir and Geils
2002). However, the long incubation period
before the development of aerial shoots
makes it initially difficult to detect the para-
site. These hidden infections are referred to
as “latent infections”—infections that have
not yet produced visible mistletoe shoots.
They complicate control efforts consider-
ably, as it is nearly impossible to eliminate all
potentially infected trees in one operational
were least common in trees ⱕ2 ft tall (5.9%
infected) and that latent infections were pos-
itively correlated with tree height at the time
of release. This finding is consistent with
Wicker and Shaw’s (1967) conclusion that
dwarf mistletoe infection in young trees is
generally low because of their smaller target
area for the dwarf mistletoe seed. Korstian
and Long (1922) were perhaps the first to
propose the idea of clearcutting and regen-
erating areas severely infested with SWDM.
Early mortality studies on the Mescalero
(Hawksworth and Lusher 1956) indicated
that many commercial stands would not
eliminate the potential for new infections
from the ⬎2 ft tall component, and release
the seedlings from suppression.
Several projects were implemented at
Mescalero over the past two decades based
on this recommendation. Tribal conscious-
ness of the severity of the mistletoe problem
is largely responsible for the high degree of
active management. Treatments designed to
control mistletoe on potentially commercial
sites should enable these stands to produce
fiber in support of the local rural economy.
Because of the high incidence and severity of
the pathogen, these treatments designed to

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entry without compromising stocking re-
quirements. Typically, the latent period is
2– 6 years, but this varies for different dwarf
mistletoe species (Hawksworth and Wiens
1996).
Mescalero’s severe SWDM infestation
is largely the legacy of past management
practices (Shields 1953, Conklin 2000). Be-
ginning with the arrival of European settlers,
overgrazing and fire suppression altered for-
est conditions and enabled a proliferation of
pine regeneration. Isolated mistletoe infec-
tions present in the mature overstory were
then able to disperse downward, dramati-
cally increasing the incidence and severity of
infection in the newly established cohort
(Shields 1953). Early foresters at Mescalero
recognized the severity of the dwarf mistle-
toe infestation and initiated control efforts.
However, their operations did not take into
account the epidemiology of the parasite,
and they were unsuccessful at controlling
SWDM. The partial cutting methods used
increased sunlight to retained trees, stimu-
lated latent infections, and actually favored
the build-up of mistletoe on many sites
(Conklin 2000). Hawksworth et al. (1977)
concluded that a century of experience has
demonstrated it is virtually impossible to
eliminate dwarf mistletoe populations
through partial cutting because of latent
infections.
Early studies on the control of SWDM
at the Fort Valley Experimental Forest near
Flagstaff, Arizona (Gill and Hawksworth
1954) indicated a prolonged latent period
after treatment. Hawksworth (1961) inves-
tigated and concluded that latency ranged
from 2– 8 years with a mean of 5 years for
this mistletoe. Most aerial shoots (92%)
were produced 3–5 years after initial infec-
tion (Hawksworth 1961). More recently,
Conklin (2002) studied the emergence of
SWDM plants from latent infections at
Mescalero and showed that these infections
reach maturity because of the high incidence mitigate the disease without compromising
and severity of the pathogen. Because of the stocking are generally not viewed as overly
unpredictable, arid, and fire-prone nature of aggressive by tribal members.
ponderosa pine sites in the Southwest, arti- To evaluate the effectiveness of their
ficial regeneration has been employed with current recommendations for treating mod-
limited success. Therefore, refinement of sil- erate to severe SWDM infestations, the
vicultural practices was required before ap- Mescalero Bureau of Indian Affairs estab-
propriate landscape-scale mistletoe treat- lished monitoring plots after an implemen-
ments could be implemented without tation. This provided an opportunity to ob-
compromising sufficient stocking levels. Be- serve the incidence of retained ponderosa
cause no viable chemical or biological con- pine with latent infections (Figure 1). The
trols are currently available for treating anticipation was that the incidence of latent
stands (Beatty and Mathiasen 2003), the infections in the seedlings would be low and
only practical control for dwarf mistletoes could be reduced even further with fol-
over large forested areas is through silvicul- low-up sanitation cutting. Results from 9
tural manipulation (Geils et al. 2002). years of monitoring are presented in this
The current recommendation for treat- article.
ing moderate to severe SWDM infestations
at Mescalero is to temporarily implement Methods
even-aged management. Operations include
a commercial harvest where all host trees Site Description
ⱖ9-in. dbh and greater are removed, fol- The 460,414-acre Mescalero Apache
lowed by a stand improvement/sanitation Indian Reservation is located in the Sacra-
cutting that slashes all remaining host trees mento and White Mountains of southcen-
⬎2 ft tall. The intent of the treatment is to tral New Mexico. Local climatic factors
retain a high percentage of uninfected trees, range from an average minimum tempera-
Management and Policy Implications
Temporary utilization of an even-aged management approach may be needed to improve the health and
vigor of some ponderosa pine stands severely infested with dwarf mistletoe. The objective of the treatment
presented in this article was to efficiently and cost effectively improve future timber merchantability
through the use of an innovative strategy. Although this treatment will probably be viewed as too
aggressive to meet many other land management objectives, it may be necessary to treat severely infested
stands if the infected species is to remain as a viable stand component. In the past, improvement of
severely infested stands has required substantial costs associated with site preparation and reforestation
planting. Now, given adequate stocking of seedlings ⱕ2 ft tall within a proposed treatment area,
mistletoe incidence and severity can be minimized in a single entry and can nearly be eradicated with
well-timed follow-up noncommercial treatments. To address forest health concerns and decrease the risk
of catastrophic wildfire, treatment options for stands severely infested with southwestern dwarf mistletoe
are a necessity. Our study demonstrated that this approach can be used to control the native parasitic plant
in the Sacramento Mountains.

380 Journal of Forestry • September 2017


Figure 1. Southwestern dwarf mistletoe (Arceuthobium vaginatum subsp. cryptopodum) low
on the bole of a ponderosa pine (Pinus ponderosa var. scopulorum) sapling at Mescalero
(black tape marks 1 ft from the ground), fall 2015.
Figure 2. Forest managers at Mescalero conducting plot re-measurement in 2012.
ture of 32.1° F to an average maximum tem-
perature of 65.9° F. The elevation ranges be-
tween 6,500 and 12,000 ft. Average annual
precipitation varies from about 15 in. at
lower elevations to ⬎32 in. at higher eleva-
tions with a majority of the precipitation oc-
curring from June to August. Forest cover is
composed of both timber and woodland
species. The managed forest area at Mes-
calero is approximately 185,000 acres, con-
sisting predominantly of ponderosa pine
and mixed conifer.
Treatment
Before the treatment, 172 0.01-acre cir-
cular plots were installed in the proposed
project area on a 330 ⫻ 330 ft grid to inven-
tory ponderosa pine regeneration. Results
indicated that ample stocking (⬎200 trees/
acre) of natural ponderosa pine ⱕ2 ft tall
was universally present.
In 2005, a commercial harvest occurred
on 7,500 acres. This included removal of all
ponderosa pine ⱖ9-in. dbh on approxi-
mately 3,300 acres where dwarf mistletoe in-
festation was severe. Feller-bunchers were
used to cut and pile merchantable timber,
and skidders were restricted to designated
skid trails to limit damage to the existing
ponderosa pine regeneration. Healthy crop
quality Douglas-fir (Pseudotsuga menziesii
[Mirbel] Franco) and southwestern white
pine (Pinus strobiformis Engelmann) were
maintained on the site. Stand improvement/
sanitation cutting operations began in fall
2005 and concluded in spring 2006. They
consisted of slashing all remaining pon-
derosa pine trees ⬎2 ft tall on the 3,300
acres.
In 2008, 12 fixed-radius plots were ran-
domly established in an interior portion of
the treatment area. Initially, seven 0.2-acre
plots were established; however, plot size
was reduced to 0.1 acre for the remaining
five plots due to high stocking. All pon-
derosa pine on each plot determined to have
been ⬎2 ft tall during the stand improve-
ment/sanitation cutting in 2006 were
slashed in 2008. This was accomplished by
measuring tree height to the top of the 2006
growth. All trees visibly infected with mistle-
toe were cut and tallied at the time of plot
installation. The remaining trees were in-
spected for mistletoe plants, distance and
azimuth from plot center were recorded, tree
heights were measured to the nearest half
foot, identification tags were placed on re-
tained trees, and crop tree quality (based on
growth form and apparent vigor) was esti-
mated. Both crop and noncrop trees were
retained throughout this study.
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Plots were first remeasured in 2012
(Figure 2). At this time, all trees were in-
spected for visible SWDM infection, height
was measured to the nearest half of a foot, all
infected trees were destroyed, and tree qual-
ity (crop or noncrop) was again noted. Seed-
lings not present in 2008 were tagged and
incorporated into the study population. In
2015, the plots were remeasured using the
same protocol as the 2012 measurement.
The plot data were then organized to pro-
duce summary tables.
Results
Based on the 2015 measurement, the
average stocking of ponderosa pine across
the project area was 1,058 trees/acre. Addi-
tional tree species present included Douglas-
fir, southwestern white pine, juniper (Juni-
perus spp.), and oak (Quercus spp.) that
make up a small component of the develop-
ing ponderosa pine stands in this geographic
area. Of the ponderosa pine stems present,
95% were determined to be of crop tree
quality. Mean tree height was 4.3 ft with a
median of 4 ft and a range of 0.5 to 11 ft
(Figure 3).
A total of 51 (2.3%) of the sample trees
had become visibly infected by 2015, 9 years
posttreatment (Table 1). This number in-
cluded trees that were infected when the
plots were established in 2008 and all addi-
tional infections found during the 2012 and
2015 remeasurements (Table 2). This per-
centage was based on a sample size of 2,263
trees, which was the sum of all the live trees
in 2015, the trees that had died from 2008
to 2015, and the infected trees that were cut
during each measurement (Table 2).
None of the SWDM plants detected
from 2008 to 2015 originated on ponderosa
pine tissue that appeared to have developed
after 2006. Because recent tissue (⬍5 years
old) is more susceptible to infection than
older tissue (Hawksworth 1961, Hawk-
Journal of Forestry • September 2017 381
 with observed latent infections (24 of 251
trees, 9.6%), which was 3 times higher than
that for the other plots (Table 1). Plot 11
had almost 5% of the sampled trees with
latent infections; however, the sample size
for plot 11 (44 trees) was relatively small be-
cause this plot was only 0.1 acre. The re-
mainder of the 0.01-acre plots (8, 9, 10, and
12) had no trees express latent infections.
Plot size influenced the number of trees sam-
pled, and this was correlated with the num-
ber of latent infections expressed on each
plot. However, plots 2 and 12 had no trees
express latent infections during the 9-year

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study, and they had ⬎90 trees sampled (143
and 95, respectively).
Previous studies of ponderosa pine and
other conifers demonstrated that the inci-
dence and rate of infection of new regenera-
tion were influenced by the severity of the
infection in the overstory, the density of re-
generation, and the age or tree height of the
regeneration (Hawksworth 1961, Scharpf
1969, Hawksworth et al. 1977, Parmeter
1978, Mathiasen 1998). The general patch-
iness of mistletoe infestation on both the
landscape and stand scale may have been
partly responsible for the plot-by-plot differ-
Figure 3. A. Stand improvement/sanitation cutting operations 2006, retained pine seed- ence observed in the incidence of trees
lings have been painted blue (trees on the ridge top were eventually masticated). B. with latent infections (Hawksworth 1961,
Posttreatment stand in 2015. Hawksworth and Wiens 1996, Conklin
2000). Although SWDM infestations in
Table 1. Plot size, number of sample trees, Table 2. Number of live trees, mortality, ponderosa pine stands on the Mescalero are
number of trees with latent infections, and and cut mistletoe-infected trees for each often severe, the severity and size of the in-
incidence of trees with latent infections by plot sampled in the study area from 2008 festations vary across these forests (Hawk-
plot for all sampling entries in study area. to 2015. sworth and Lusher 1956).
More than 98% of the pretreatment re-
Incidence of Live generation survey plots had infected over-
trees with Plot trees Mortality Mistletoe cut story trees within a distance that could have
Trees with latent no. 2015 2008–2015 2008 2012 2015 Total infected trees on the plot (35 ft). However,
Plot Plot Sample latent infections
no. size (ac) trees infection (%) 1 95 23 5 1 1 125 this is the extent of pretreatment infection
2 137 6 0 0 0 143 data, and we were unable to definitively
1 0.2 125 7 5.6 3 476 44 3 0 4 527 link the relatively high incidence of trees
2 0.2 143 0 0 4 74 19 1 0 0 94
3 0.2 527 7 1.3 5 441 21 2 1 2 467 with latent infections observed on plot 6
4 0.2 94 1 1.1 6 200 27 15 6 3 251 (9.6%) and relatively low incidence on
5 0.2 467 5 1.1 7 348 17 0 1 4 370 other plots with pretreatment overstory
6 0.2 251 24 9.6 8 40 3 0 0 0 43
7 0.2 370 5 1.4 9 49 5 0 0 0 54
infection incidence.
8 0.1 43 0 0 10 47 3 0 0 0 50 Total plot mortality consisted of 202
9 0.1 54 0 0 11 39 3 2 0 0 44 dead trees (9%) from 2008 to 2015. The
10 0.1 50 0 0 12 64 31 0 0 0 95 majority of the mortality (80%) occurred
11 0.1 44 2 4.5 Total 2010 202 28 9 14 2,263
12 0.1 95 0 0 between 2008 and 2012. Rodent girdling
Total 1.9 2,263 51 2.3 and drought were suspected to be the pri-
in 2008 2 years after the treatment, 9 in mary drivers of mortality from 2008 to
2012, and 14 in 2015 (Table 2). 2012. Historical analysis of the Palmer
sworth and Wiens 1996), if any new infec- The highest incidence of latent infec- drought severity index (PDSI) for Mescalero
tions occurred, they would probably have tions occurred on trees in plots 1 and 6 over the study period indicated that the first
appeared on the younger host tissue. where between 5 and 10% of the sample protracted extreme drought conditions dur-
Twenty-eight trees with latent infections trees were infected before the 2006 treat- ing the study occurred during 2011 (Na-
were found at the time of plot installation ment; plot 6 had the greatest number of trees tional Oceanic and Atmospheric Adminis-

382 Journal of Forestry • September 2017

tration 2015). The mortality that occurred
from 2012 to 2015 (20% of the total) was
observationally attributed to elk (Cervus ela-
phus subsp. nelsoni Bailey) rub damage. The
elk herd at Mescalero was ⬎4,200 head with
72% of the cows giving birth every year
(Mescalero Apache Tribe 2015).

Discussion
Two years after treatment, the majority
of ponderosa pines with latent infections
that were initially ⱕ2 ft tall showed signs of
infection. After 6 years, ⬎70% of the trees
with latent infections were observed, indi-

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cating that an additional sanitation cutting
conducted 5 years after initial stand im-
provement/sanitation cutting would have
detected and removed the majority of the
infected regeneration. Risk of catastrophic
wildfire persisted after treatment; however,
this was likely reduced once the majority of
trees reached breast height. Ponderosa pine
is generally regarded as being resilient to
ground fire once it reaches 4.5 ft tall (Bailey
and Covington 2002).
The number of trees with latent infec-
tions detected 9 years after the treatment (14
trees) was larger than that 6 years after the
treatment (9 trees); however, it is probable,
based on artificial inoculation experiments
conducted in Arizona (Hawksworth 1961),
that nearly all of the latent infections have
been observed. Another remeasurement is
needed to estimate the incidence of trees that
express latent infections after 9 years and the
long-term impacts of this treatment. It
should be noted that although Hawksworth
(1961) reported that 8 years was the maxi-
mum incubation period based on his obser-
vations of artificially inoculated ponderosa
pines, this study and that of Conklin (2002)
indicated that a longer incubation period of
at least 9 years may occur at Mescalero.
Host susceptibility to dwarf mistletoe is
rated using a subjective classification system
based on the percentage of trees of the host
species in question that are infected by dwarf
mistletoe within 19.7 ft of a principal host
heavily infected with the same species of
dwarf mistletoe (Geils et al. 2002). Accord-
ing to these standards, ponderosa pine is
considered a principal host (⬎90% in-
fected) of SWDM. Thus, the resulting low
incidence (2.3%) of trees with latent infec-
tions observed is encouraging.
Regeneration surveys over many de-
cades suggest that regeneration of ponderosa
pine at Mescalero is more reliable than on
many sites in the Southwest. The survival
Figure 4. Ponderosa pine (Pinus ponderosa var. scopulorum) seedling at Mescalero that
apparently sprouted from epicormic buds below stump height, summer 2015.
and growth of regeneration in this study fit
that trend with current stocking conditions
more than 1 order of magnitude higher than
reference conditions for the region (Fulé
et al. 1997). Although we anticipate signifi-
cant mortality because of competition in the
future, it should be noted that all of the mor-
tality that occurred on the plots from 2006
to 2015 was attributed to either drought or
animal damage and not to obvious SWDM
infection.
Many ponderosa pines slashed on the
plots during the 2005 to 2006 treatment
sprouted from epicormic buds below the cut
(Figure 4), increasing the stocking levels es-
timated from the pretreatment surveys.
Sprouting is a natural physiological reaction
of plants to disturbance that probably devel-
oped as a survival mechanism to either fire or
flooding (Baker 2009). Although many tree
species (including some pines) are known to
sprout from their original root stock after
disturbance, to our knowledge, this behavior
has never been documented in Pinus pon-
derosa var. scopulorum. Nevertheless, 129
(5.7%) ponderosa pine trees in this study
apparently sprouted after slashing. This re-
sponse from ponderosa pine may be associ-
ated with a genetic predisposition, high site
quality, stump size, and/or the seasonality of
treatments. Further investigation is needed
to definitively document the phenomenon.
One of the infected trees removed from
plot 7 in 2015 was a sprout. Dwarf mistletoe
aerial shoots appeared both below and above
where the tree was cut, indicating that an
infection existed below stump height (ap-
proximately 4 in.) and grew a short distance
into the new sprout. Therefore, trees that
sprout after treatment may be more likely to
be infected because the original cut tree was
a larger target for mistletoe seed, and the par-
asite may have established below stump
height.
Conclusions
The observations from this study sup-
port the management recommendations of
Hawksworth (1961) which state that regen-
eration within SWDM-infested ponderosa
pine stands should be retreated with sanita-
tion cutting at recurrent 5-year intervals.
Our findings confirm Wicker and Shaw’s
(1967) conclusion that dwarf mistletoe in-
fection in young trees is generally low be-
cause of their smaller target area and endorse
the observation of Conklin (2002) that
SWDM latent infections were minimal in
trees ⱕ2 ft tall. Because of the observational

Journal of Forestry • September 2017 383

nature of this study, we can draw only lim-
ited conclusions from its results.
A prolonged incubation period is the
primary reason that complete eradication of
SWDM is impractical in most settings. Our
observations demonstrate that implementa-
tion of the locally developed management
guidelines for moderate to severely infected
stands resulted in a low incidence of latent
infections (⬍3%) in the residual cohort and
were used to control SWDM on commercial
ponderosa pine sites in the Sacramento
Mountains. This management technique
may be applicable in other SWDM infested
pine stands because of the epidemiology of
SWDM related to the dispersal of seed and
infection of regeneration.
Treatments that cut all hosts ⬎2 ft tall
followed by sanitation cutting 5 and 10 years
after the initial stand entry may be the most
economically efficient way to control
SWDM in severely infested pine stands.
This treatment requires sufficient natural re-
generation ⱕ2 ft tall before implementa-
tion. In certain areas of the Southwest, re-
generation is sporadic; therefore, treatments
would need to be designed to capitalize on
these events. Future observations will pro-
vide more detailed data on the total num-
ber of retreatments required to eliminate
SWDM latent infections in developing
stands.
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