Potential Mountain Pine Beetle (Coleoptera: Scolytidae)

Attack of Lodgepole Pine as Described by

Stand Density Index

JOHN A. ANHOLD AND MICHAEL J. JENKINS

Department of Forest Resources, Utah State University,
Logan, Utah 84322-5215

Environ. Entomol. 16: 738-742 (1987)

ABSTRACT Ninety-four unmanaged lodgepole pine stands, from a broad geographical
range in the western United States, were examined to evaluate the relationship between
stand density and susceptibility to mountain pine beetle (MPB), Dendroctonu8 pondero8ae
Hopkins, attack. Population trends were not significantly related to variation in stand density
as measured by stand density index (SDI). Percentage of trees killed per hectare by MPB in
stands with >80% lodgepole pine did vary significantly with changes in SDI. From these
data three SDI zones were identified as follows: 1) stands with SDI's of <125 showed low
potential for attack, 2) stands between 125 and 250 SDI showed much greater levels of tree
mortality, gradually decreasing toward the 250 SDI, 3) tree mortality decreased in stands as
density increased beyond the 250 SDI value.

KEY WORDS Dendroctonu8 ponderosae, lodgepole pine, stand density index

THE MOUNTAINPINEbeetle (MPB), Dendroctonus
ponderosae Hopkins, represents the definition of
its genus: killer of trees. Large-diameter, thick-
phloemed lodgepole pine (LPP), Pinus contorta
Douglas var. latifalia Engelmann, are mass at-
tacked, accelerating epidemics as the MPB period-
ically reaches outbreak numbers across the western
United States and Canada (Amman 1972, Berry-
man 1976, Klein et al. 1980). Once an outbreak is
established, there seems to be no rigorous pattern
by which attacks on stands occur. The population
density of MPB apparently determines the number
of trees that will be successfully attacked (i.e., more
beetles are required to overcome the resinosis de-
fenses of large, vigorous trees compared with small-
er, weaker trees [Raffa & Berryman 1983]).
Adult MPB attack LPP in July or August, intro-
ducing blue-stain fungi while feeding on and con-
structing egg galleries in the phloem layer (Amman
1978). Eggs hatch and larvae feed on the phloem,
girdling and killing the tree, aided by blue-stain
fungi (Amman 1978, Ballard et al. 1984). Larvae
overwinter, completing the life cycle in the spring.
Investigations of stand/site conditions indicate
that losses to MPB can be minimized through sil-
vicultural stand treatments (Hamel 1978, Waring
& Pitman 1985). Cole (1978), however, pointed out
the need for identifying an accurate risk classifi-
cation scheme for LPP stands to assess susceptibility
to MPB. Several MPB risk classification systems
have been developed (Mahoney 1978, Amman
1983); however, these systems need to be evaluated
to determine their validity, accuracy, and utility
over broad geographic areas.
Stand density index (Sm) was examined by Stuart
(1984) as a predictor of MPB outbreaks in climax
Pinus contorta var. murrayana Grev. & Balf. stands
in south-central Oregon. This study was designed
to examine SDI further in LPP var. latifolia over
an extensive geographic area as a simple means of
evaluating stand susceptibility. If SDI correlates
as a predictor of susceptibility, then ~cCarter &
Long's (1986) LPP density management diagram
could be used as a risk rating system.
Materials and Methods
In the summer of 1984, 300 stands were exam-
ined throughout the western United States (Wash-
ington, Oregon, Montana, Wyoming, Utah, and
Colorado) (Fig. 1). Suitable candidate stands were
selected at random throughout the study area using
the following criteria: 1) high climatic suitability
(i.e., low elevation, moderate winter temperatures);
2) no recent disturbance by man or any high degree
of natural disturbance, wind, fire, etc.; 3) 25% or
more LPP; and 4) uniformity of habitat type, stand
structure, species composition, and canopy config-
uration within each stand. Stands ranged in infes-
tation history from no recent infestation to those
stands currently undergoing an outbreak. Under
these conditions, stands were considered climati-
cally, geographically, and otherwise suitable for
MPB establishment.
Each stand was sampled by establishing 10 plots
per stand, located 100 m apart using two lines (five
plots per line). Some modifications of plot location
were made as dictated by stand shape, but a 100-m
spacing was maintained. After randomly locating
plot center, data were collected as follows: I) di-
ameter and species (12.7 cm diameter at breast
height [dbh] and larger); 2) alive or dead (cause of

738
June 1987 ANHOLD & JENKINS:STANDDENSITYINDEX ASAN INDICATOR 739

,.
120 110 100 90 Table l. Average characteristics of stands used to plot

, .
,,
MPH attack rate data on Fig. 2

No.of Avgphloem
, , I
, , I
• I I Attackrate stands thickness Dq SDI
" : ~ ,,' (mm)
50 "I : ~ /
Increasing 25 1.24 9.0 191
'.... I • '
\
,
I
I
I
I
'
\ Decreasing 30 1.75 8.5 238
": l ~
Static 39 2.06 8.4 198
--_......
•"::i,j ----- ::
,.__ ~
..i __ -.,
I
- .••.••• .••

_____
, ~...t
: ••~::
: I
"
"

:
) ~, .:-
''--:- .. _----_.~
~--- -- --J..
: was developed were used (i.e., even-aged LPP

40
-...•
:1: _
--'--_4__ 1-.
I :'"
:
,-------
r---
.••••
,
stands where 80% of the total basal area is LPP).
Also, stands had to have MPB activity.
: r--i..: \!;-
:
'..
",
: 4.-,-"'C------.
:
I
:
I
~...
q
;---------
,
\..
Percentage of the total TPH attacked by MPB
was determined for each stand and plotted against
its respective SDI. Zones of SDI were determined
'\ I : :

',\, ....
!-------t- -..-- - -(':_-_:.-
..-- -- on the basis of stand growth and development and
\~~ : I: overlaid on the plot. Zone A had an SDI from 0 to
')
, " :I '-"-""-1" 125, zone B from 126 to 250, and zone Chad SDI
S : : values >250. An SDI of 125 corresponded with
30 ......
.••• __~_r-
I .J:.--""-'
'\" crown closure and the onset of competitive inter-
''\ action, and an SDI of 250 represented the begin-
\ .••.
,~
,...- .•..•.•., ning of full site occupancy (McCarter & Long 1986).
'.'. Average phloem thicknesses were calculated for
'- each of the stands and plotted against SDI.
The plotted data were first visually analyzed for
Fig. 1. Approximate location of study sites in the signs of grouping among MPB attack rate values
western United States.Each symbol represents fivestands. and within zones. One-way analysis of variance was
conducted to determine if any significant differ-
death); 3) year tree attacked by MPB: a) current ences existed. The t test with one degree of free-
year, b) last year's attack, c) 2 yr and older attacks; dom was used to test for specific differences in TPH
4) unsuccessful and strip attacks (current or old); attacked in zones A, B, and C, at the 95% confi-
5) phloem thickness (green LPP only); two samples, dence level.
180 apart, at breast height. Squares (ca. 1.27 cm)
0

were removed after four cuts had been made using Results
a sharp hatchet. The sample was then visually cal-
ibrated with a ruler graduated in millimeters. Ninety-four stands met the species composition
Compression of phloem was minimized by using criterion as outlined for McCarter & Long's (1986)
sharp tools. diagram (Fig. 2). The SDI levels ranged from 92
SDI (Reineke 1933) values, which express the to 492, with Dq's ranging from 16.5 to 33.8 cm.
relation of trees per hectare (TPH) and quadratic Table 1 shows the average phloem thickness, Dq,
mean diameter (Dq) (Sm = TPH(Dq/25)L6, were SDI, and number of stands in each category. It was
calculated for each stand along with a value de- expected that the increasing stands would have a
termined for each stand from the number of cur- higher sm than the decreasing stands; however,
rent to previous year's attacked trees. An increasing just the opposite occurred. No correlation existed
MPB attack rate coincided with a stand having a between the beetle attack rate values and SDI, Dq,
larger number of TPH currently attacked (includ- or phloem thickness.
ing current and current strip attacks) than previous Plotting sm against percent TPH attacked for
year's TPH attacked. The reverse would be true the same stands revealed a left-skewed distribution
for stand with a decreasing MPB attack rate. Stands (Fig. 3). Low mortality was observed below the
with equal numbers of current and previous year's 150 SDI level. At that point a sharp increase in
attacks were considered to have a static MPB attack percent TPH attacked occurred, followed by a
rate. Admittedly, a tighter trend could be formu- gradual tapering off with increasing SDI. A sig-
lated if older attacks were used. However, deter- nificant difference was found in TPH attacked be-
mining year of attack beyond 2 yr is not accurate; tween zones A and Band Band C at the 95%
thus, attacks older than 2 yr were grouped as older confidence level.
attacks. Average phloem thickness and sm for each stand
The individual stands were plotted on McCarter were plotted, indicating a random distribution.
& Long's (1986) LPP density management dia- However, after eliminating stands with >30% TPH
gram using calculated TPH and Dq. However, only attack a pattern appeared (Fig. 4). Stands with high
stands that met the criterion on which the diagram attack percentages do not show representive av-
740 ENVIRONMENTAL ENTOMOLOGY Vol. 16, no. 3

700
600
500
400
300
X 30.0
w
o
~ 25.0 M PB Attack Rate
)- 200
t- • INCREASING
~ \50-20.0
w Q STATIC
o
~ \00 15.0
* DECREASING

t!
(/)

50-10.0
9.0
8.0

"; 5.0
cu
s;
CJ
C
-= 4.0
Q:
W
t-
W
~ 3.0
«
C

2.0

1.0
50 100 1000

TREES PER ACRE

Fig. 2. Ninety-four LPP stands, depicting increasing, static, and decreasing MPB attack rate values, plotted on
the LPP density management diagram (McCarter & Long 1986).
June 1987 ANHOLD & JENKINS:STANDDENSITYINDEX ASAN INDICATOR 741

100 3.0 Zen. C

Zone A ZoneS
Zon •• Zon. a Zon. C 0
90 0 0 0 o 0 0
2.5 It 0 ~oo 0 8
eo 0 o 000
0 00 0 0 0
70 0 0
o 00
8
"0
Q)
..>t.
•• E
E
2.0
00
6> 0
60
0
1l 0
E
<: so
0
0 0 Q)
0
1.5 0
:c 0 0
~ -40 o 0 0
0
0- 00
0
0 0
0 0 0
~ 0 til
~ 1.0 0 0
30 00 0
80 0
0
20 0 00 0
0 0.5
o 930 0
10 SOO <9 0
o 0
o fi,0 00
0% 0
o 0 0
000
0 0
50 100 150 200 250 300 350 400 450 50 100 150 200 250 300 350 400 450
Stand Density Index Stand Density Index
Fig. 3. Potential LPP losses to MPB for' different Fig. 4. Relationship of SDI and LPP phloem thick-
SDI's. ness.

erages because larger, thicker-phloemed trees are secondary resin formation (Shrimpton 1978). It is
selected and killed. Stands have a stable phloem also possible that a physiological change within the
thickness until an SDI of ca. 225 followed by a trees has lowered the resistance to fungal inocu-
decline with increasing SDI. lations vectored by MPB (Raffa & Berryman 1982).
A reduction in the defense mechanism, while am-
ple food supplies are present, predisposes a stand
Discussion
to potentially high levels of attack. These processes
Phloem thickness has long been implicated as a become significant at an SDI of ca. 150, shortly
causal agent of MPB infestations (Amman 1969). after crown closure. It should be noted that SDI
The rationale is that temporarily stressed, thick- values were determined using trees ~12.7 cm in
phloem trees are predisposed to MPB attack and diameter. If trees of smaller diameter are included,
produce greater brood numbers because of the the SDI levels increase. This may cause the critical
thicker phloem (Berryman 1982). Stress also re- 150 sm to shift to a higher sm.
duces resin flow, adversely affecting the tree's ma- In unmanaged stands, irregular spacing is com-
jor defense mechanism, resinosis. Factors that dis- mon, resulting in uneven competition throughout
rupt or impair functioning of the resin system a stand, with total competitive interaction occur-
influence host susceptibility (e.g., moisture stress or ring after an SDI of 125 is reached. This may
fungal establishment within the xylem). Increasing explain the delay in high levels of attack after a
stand density may cause moisture stress, hence in- stand enters zone B.
creasing host susceptibility. Increased susceptibility As a stand continues through zones Band C, the
has been shown to be associated with reduced radial maximum growth potential of individual trees con-
growth (Mahoney 1978) and thicker phloem (Cole tinues to decrease, phloem retention is gradually
1973). diminished (Cabrera 1978), and the potential for
This study suggests that the potential for MPB high levels of attack is lowered. Resin flow is less
attack in LPP stands goes through three phases. than its maximum potential, but because of the
First, a period of low potential mortality, followed thinner phloem MPB are not as likely to attack.
by a phase of high attack potential, and finally a However, if MPB populations are large enough,
period of declining probability of attack (Fig. 3). trees can be attacked regardless of phloem thick-
The first phase, zone A, represents fast-growing, ness, although brood production decreases with de-
noncompeting trees, growing at their maximum creasing phloem thickness.
potential for the given site conditions. Resin pro- Full site occupancy is reached at an SDI of 250.
duction and flow should be at high levels, account- This is of importance from a managerial standpoint
ing for the low percentage of TPH attacked. As (i.e., if wood production is the major objective, any
stand growth continues, leaving zone A, competi- growth below an SDI of 250 results in a loss of
tive interaction begins. At an SDI of ca. 125, the potential stand growth). The SDI boundary be-
growth rate of individual trees is something less tween 400 and 700 is the zone of imminent com-
than their maximum potential. Some level of stress petition mortality (Drew & Flewelling 1979,521),
has been initiated with the transition from zone A representing the lower and upper limits of self-
to B. At this point phloem thickness is suitable for thinning for LPP. Lower attack and brood pro-
brood production (Amman 1972), but low levels of duction rates are expected because of thinner
tree stress are causing declines in resin flow and phloem (Fig. 3 and 4).
742 ENVIRONMENTAL
This study indicated that regardless of the SDI,
within the ranges examined, MPB activity can be
expected to increase, decrease, or remain static (Fig.
3). This was not expected; rather, grouping of MPB
attack rate values at separate SDI ranges seemed
more intuitive. It was hypothesized that stands with
increasing values would coincide with those of less
vigor, higher competitive interaction, and, thus,
higher SDI ranges. Area-wide MPB population
levels are very important in determining attack
rates in any stand with suitable SDI and phloem
thickness.
Acknowledgment
The help of David Holland, James Long, James
McCarter, and Mark McGregor is gratefully acknowl-
edged. We appreciate the assistance of Gene Amman in
obtaining data from the USDA Forest Service Canada/
United States Mountain Pine Beetle Lodgepole Pine Haz-
ard Rating project. This is Journal Paper No. 3265 of the
Utah Agricultural Experiment Station.
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Received for publication 27 May 1986; accepted 18
February 1987.