------·
Estimating Pre-treatment Variation in the Oak Leaf-chewing Insect Fauna of the Missouri Ozark Forest Ecosystem Project (MOFEP)
Robert J. Marquis and J osiane Le Corffl
Abstract.-We describe spatial and temporal variation in the insect herbivore communities associated with the MOFEP, prior to applica tion of contrasting cutting regimes. No pre-treatment differences were found in total insect density on either black (Quercus velutina) or white oak (Q. alba) during 1993-1995. There was great seasonal variation in insect abundance on both host species as well as high variation across years for white oak. White oak on north- and east facing slopes tended to have more insects than white oak on south and west-facing slopes. Sites under the auspices of the Missouri Department of Conservation for a longer time had higher insect densities than more recently acquired sites.
The goal of the Missouri Forest Ecosystem levels can have negative impacts on growth and Project (MOFEP) is to determine the effects of survivorship of trees (Rexrode 1971, Coffelt et altemative forest management schemes on al. 1993) and can potentially influence ecosys long-term forest productivity, genetic structure tem processes such as nutrient ~ycling(Parker of plant populations, and biodiversity of the and Patton 1975). Th~resultant economic communities under management. Treatments impact could be substantial because timber (even-aged versus uneven-aged management, harvesting and processing in Missouri generate plus controls; see Brookshire et al. 1997 for $3 billion in economic activity annually (T. explanation of the MOFEP treatments and Robison, personal communication). Oaks are design) were begun in 1996. During 1993- also important as food for wildlife (Brezner 1995, sampling was carried out to quantify any 1972, White 1995), but their economic contri differences that might exist among replicates, so bution in this form has yet to be estimated. that potential true effects produced by the treatments could be distinguished from condi From a conservation standpoint, Lepidoptera, tions existing prior to treatment application. and butterflies in particular, have come under increasing scrutiny as the abundance of many One of the major components of these Ozark species declines (e.g., Thomas and Mallorie forested communities, in terms of their ecologi 1985, Hill1995, Legge et al. 1996). Lepidoptera cal, economic, and conservation role, are the can be used as indicator species for changes in insect herbivores that feed on oaks. The major habitat quality because their populations are ity of leaf-feeding insects on Missouri oaks are intimately tied to the abundance of their host larvae of Lepidoptera (moths and butterflies) plant, and host plant decimation is often (Marquis and Whelan 1994). As herbivores, synonomous with habitat destruction (e.g., they have potential gro\\1:h impact on their Eberhardt and Thomas 1991, Sparrow et al. hosts. Non-outbreak population levels cause 1994). In Missouri, these insects represent a enough damage to reduce growth of saplings major component of the State's biodiversity and (Marquis and Whelan 1994), while outbreak natural heritage: at least 300 species have been documented on oaks in the Eastem United States and 200 on white oak alone in Missouri (Tietz 1972, Covell 1984). Additionally, these 1Associate Professor of Biology and Postdoctoral insects are important because they serve as Research Associate, respectively, Department of hosts for a number of parasitoids, both insects Biology, University of Missouri-St. Louis, 8001 and nematodes. In so doing, they could serve to Natural Bridge Road, St. Louis, MO 63121- maintain a reservoir population of parasitoids 4499. that might be used to control invading exotic 332 insect pests such as the European gypsy moth, pattems. Results of analyses of herbivore Lymantria dispar. community composition and canopy sampling will be presented separately. The goal of this project was to describe variation in the abundance of the leaf-chewing insect METHODS fauna among replicate sites of the MOFEP project during the 3 years before ha.IVesting General Census Methods treatments were applied. Two host plant spe cies were chosen for study, black (Quercus To determine insect abundance, leaves were veZutina) and white (Q. alba) oaks. These two searched both top and bottom, as were associ species dominate the canopy of most MOFEP ated branches and the main stem of the tree. upland stands. Limited resources prevented us Chewing insects encountered were classified to from sampling more host species. We chose to morpho-species. Each census person was given include all leaf-chewing insect species, and not a training period in identification prior to actual a subset of that fauna, because preliminary sampling, and had a list of descriptions for all sampling in 1989-1992 suggested that species' morpho-species known to be encountered. At abundances of leaf-chewing insects fluctuate no time were leaves collected. All insects were unpredictably (RJM, unpublished data). Thus, left intact on the plant unless individuals were a study that focused on only common species in unknown. In that case, unknowns were taken one year would have no estimates of abun back to the laboratory for rearing and photo dances of previously rare species that had graphing. Each unknown was given its own become common. High diversity of the leaf unique sample number and description, and chewing guild (at least 200 species) prevented this information was entered into a database. us from sampling other herbivore guilds associ Photographs were taken, and the insect was ated with black and white oak. However, num observed through development until it could be bers of galling and sap-sucking insects in the verified as a previously-recognized species or MOFEP region, and throughout Missouri, classified as a species new to our inventory. appear to be low (Marquis and Whelan 1994). Photos were used in the field to help with identification when necessary. Our overall objective here is to describe pre treatment variation (years 1993-1995) in the Preliminary Sampling to Establish abundance of the leaf-chewing insect herbivores Adequate Sample Size associated with Quercus alba and Q. veZutina at the MOFEP sites. In this paper, we first de Preliminary sampling was conducted in early scribe the initial sampling undertaken to estab September 1992 to determine the sample size lish the sample size chosen for the 3 years of necessary to adequately estimate the mean and pre-treatment sampling. Second, we analyze variance in total numbers of insects per tree per the pre-treatment data to determine whether stand and per site, and species composition per sites assigned to different treatments actually site. Ten trees per stand were sampled in site 6 demonstrated differences before treatments for each of three north- facing slopes and three were applied. Third, we present analysis of the south-facing slopes for both black and white spatial variation in herbivore abundances oak. The mean and variance in total numbers irrespective of the original sampling design to of insects per leaf area per tree were plotted for understand possible sources of background each stand and across stands for each tree heterogeneity among the study sites, including species. The number of new species encoun potential biases inherent in our sampling. tered with increasing number of stands sampled Fourth, we discuss whether we are currently was plotted to determine how the number of undersampling or oversampling in light of the insects species found in a site increased with pattems observed. To do so, we use power increasing number of stands sampled. The analysis to predict the minimum difference total number of species per site was compared produced by treatments that we would be able with species accumulation curves for each to establish as statistically significant. Finally, stand and each host plant. This preliminary we make recommendations about post-treat sampling then served to determine sample size ment sampling. We present results for total for all subsequent sampling in years 1993- insect numbers sampled in the understory and 1995, described below. For this analysis and all analyze abundances of a subset of species to subsequent analyses, insect abundance was illustrate the variety of individual species expressed on a leaf area basis. Average leaf 333
• •
/~ /~
years. years.
when when
When When
the the from from
Sampling Sampling nearby nearby
Stand Stand
Tree Tree
approximately approximately
Stands Stands
chosen chosen sites. sites.
cutting, cutting,
Stand Stand
The The on on
leaves leaves calland calland
sample sample three three
From From
stands stands
geographic geographic
sample. sample.
Leaf Leaf
and and
trees. trees.
3,000 3,000
hanging hanging
leaves leaves
(1993 (1993
teristics teristics leaves leaves
(ELT (ELT
conducted conducted
400 400 uneven-aged uneven-aged
assigned assigned
area area
Ground-level Ground-level
Overall Overall
sample sample
et et
334 334
al. al.
south-
~CCJ)JFJEIP ~CCJ)JFJEIP
second second
five five
MOFEP MOFEP
ha ha
and and
Attributes Attributes
per per
those those
18) 18)
stands stands
1997) 1997)
necessary necessary
1993 1993
white white
and and
To To
available, available,
Selection Selection
Sample Sample
per per
per per
per per
Dead Dead
were were
neighbors neighbors
randomly randomly
(six (six
size. size.
each. each.
MOFEP MOFEP
of of
stands stands
black black
type type
branches branches
(Sheriff (Sheriff
Leaves Leaves
tree tree
per per
Tree Tree
to to
maintain maintain
was was
and and
200 200
stand stand
tree. tree.
species, species,
1994) 1994)
from from
available available
round round
proximity proximity
to to
per per
includes includes
either either
oak oak
censused censused
design design
site. site.
individuals individuals
on on
Sampled Sampled
1993-1995 1993-1995
treatment. treatment.
species species
(ELT; (ELT;
censusing censusing
Sample Sample
Each Each
1995, 1995,
stratified stratified
oak oak
Size Size
west-facing west-facing
undamaged, undamaged,
a a
site) site)
sampled sampled
Design Design
were were
north-
we we
a a
to to
and and
0.5-
leaves leaves
distributed distributed
or or
from from
to to
of of
mixture mixture
The The
a a
of of
trees. trees.
complete complete
with with
consistency consistency
see see
June June
censused censused
(see (see
of of
were were
control, control,
we we
maintain maintain
cutting cutting
in in
nine nine
He He
sub-canopy sub-canopy
was was
to to
and and
counted counted
Size Size
these these
each each
trees trees
same same
those those
Brookshire Brookshire
and and
control control
and and
sampled sampled
within within
1.0-acre 1.0-acre
figure figure
a a
(0.5 (0.5
1997). 1997).
were were
in in
Sites Sites
census census
Censuses Censuses
selected selected
More More
estimated estimated
sites sites
maximum maximum
other other
slopes slopes of of
even-aged even-aged
east-facing east-facing
year year
fully fully
1,200 1,200
were were
(year (year
sites sites
among among
even-aged, even-aged,
the the
trees trees
to to
likely likely
saplings saplings
a a
replaced replaced
a a
either either
2 2
are are
and and
sites sites
of of
trees trees
2.5 2.5
in in
minimum minimum
general general
comparable comparable
minimum minimum
in in
area area
and and
expanded expanded
three three
(1995). (1995).
to to
spread spread
(ELT (ELT
approximately approximately
has has
2006). 2006).
timing timing
black black
and and
randomly randomly
blocked blocked
et et
to to
Brookshire Brookshire
m) m)
uneven-aged uneven-aged
five five
canopy canopy
based based
by by
were were
in in
of of
across across
al. al.
be be
per per
sites sites
been been
and and
was was
the the
stands stands
with with
17) 17)
five five
ecologi
slopes slopes
the the
white white
and and
charac
oak oak
cut cut
1997). 1997).
over over
of of
stand. stand.
of of
added added
same same
low
and and
on on
leaf leaf
by by
were were
May May
in in
a a
in in
trees). trees).
treatments treatments
replicates. replicates.
included included
interaction interaction
tested tested run run
ELT ELT
was was
to to
time time
to to year year
consistent consistent
Results Results
Brookshire Brookshire mine mine
presented presented
site site
In In subjects subjects
individual individual oak oak
common common
analyses analyses
random random
of of
temporal temporal
a a
to to
weeks weeks
August August
Whelan Whelan
leaf leaf
ses ses
the the first first
ments, ments,
For For
measures measures
per per
original original
et et among among
the the
composition composition
Due Due
always always
Number Number
split-block split-block
the the
be be
al. al.
all all
allow allow
determine determine
a a
relative relative
year: year:
3 3
(Littell (Littell
data data
effect effect
area area
for for
separately separately
separate separate
considered considered
and and
used used
to to
on on
to to
crossed crossed
north-
the the
1991, 1991,
years: years:
original original
for for
to to
-September. -September.
Because Because
changes changes
sites sites
white white
influence influence
for for censused censused
for for
insect insect
design design
statistical statistical
effect, effect,
1994), 1994),
of of
effects effects
site site
were were
herbivore herbivore
per per
collected collected
to to
degree degree
complete. complete.
effects effects
initial initial
with with
April-May, April-May,
to to
analysis analysis
as as
species species
et et
across across
block block
Samples Samples
both both
term term
across across
In In
et et
determine determine
importance importance
p. p.
6, 6,
and and
as as
test test
through through
design. design.
tree). tree).
were were
during during
al. al.
between between
repeated repeated
with with
oak oak
Statistical Statistical
al. al.
this this
experimental experimental
conducted conducted
130, 130,
density density
were were
in in
that that
5, 5,
trees trees
(Brookshire (Brookshire
they they
to to
(how (how
in in
black black
differences differences
were were
to to
repeated repeated
1991, 1991,
between between
effects effects
for for
east-facing east-facing
values values
1997. 1997.
in in
4, 4,
the the
and and
sites. sites.
pattems. pattems.
in in
be be
considered considered
or or
analysis, analysis,
comparison comparison
site, site,
herbivore herbivore
which which
of of
species species
All All
282, 282,
of of
the the
At At
3, 3,
considered considered
were were
pre-treatment pre-treatment
which which
Each Each
were were
years years
late late
among among
treatments treatments
Separate Separate
nested nested
variance variance
May May
the the
significant. significant.
subjects subjects
ELT. ELT.
black black
oak oak
if if
von von
(number (number
but but
2, 2,
measures measures
each each
and and
of of
as as
same same
The The
obtained obtained
the the
von von
year year
June, June,
1, 1,
for for
Analysis Analysis
ELT ELT
sites sites
measures measures
assigned assigned
censused censused
ELT, ELT,
census census
censuses censuses
was was
block, block,
designated designated
Ende Ende
on on
censuses censuses
1993-1995, 1993-1995,
et et
in in
7, 7,
the the
design design
ELT ELT
ELT ELT
effect effect
census, census,
goal goal
oak. oak.
slopes slopes
trees) trees)
within within
Ende Ende
the the
abundance abundance
order order
to to
al. al.
white white
and and
(Marquis (Marquis
sites sites
8, 8,
were were (AN (AN
analyses analyses
of of
effects effects
not not
census, census,
of of
to to
effects effects
late late
be be
was was
five five
was was
interacted interacted
1993) 1993)
ANOVA, ANOVA,
was was
9. 9.
1997), 1997),
any any
treated treated
for for
required required
Profile Profile
These These
insects insects
of of
OVA) OVA)
be be
site site
and and
1993) 1993) differences differences
represented represented
to to
described described
sampled, sampled,
fixed. fixed.
irrespective irrespective
treatment treatment
for for
four four
July, July,
models models
to to
and and
equal equal
ELT ELT
sites sites
most most
and and
individual individual
included included
fixed. fixed.
considered considered
to to
treat
observed observed
(effects (effects
in in
illustrate illustrate
was was
of of
were were
within within
each each
repeated repeated
and and
and and
(Littell (Littell
were were
and and
deter
same same
black black
and and
times times
analy
the the
was was
as as
was was
both both
years years
we we
per per
and and
were were
An An
2 2
with with
are are
a a
ELT ELT
run run
of of
for for
in in
of of in in Type N SS were used to calculated F- and P 5 values for effects of ELT on abundances of --+-- Mean insects on this tree species. In all other cases, -o-- Standard deviation Type III SS were used. A result was considered =~ 4 to be statisticaly significant at P = 0.05 unless = otherwise specified...... - -=~ 3 Possible site effects might occur due to seasonal ~e changes taking place within the course of a ...... 2 ....<:ll census. Because 2 weeks are required to ~ ~ complete an entire census, later censused <:ll stands and sites could have different insect ....= 1 numbers than stands and sites visited at the beginning of the spring census. Such seasonal 0~-¥~~~~~--~~~~~ changes occur most rapidly during the April o 1 2 3 4 5 6 7 s 9 ro May census, when there is a change in insect species composition and numbers associated Number of trees sampled per stand with the completion of leaf expansion (RIM and Figure example, chosen randomly from JL, unpublished data). Accordingly, the April 1.-0ne the six patterns available for white oak, May censuses were timed by phenology rather showing the relationship between the esti than calendar date to take into account the fact mate of the mean and standard deviation that leafing and insect activity vary by years. of the nwnber of insects per square meter leaf For both oak species, we tested whether species area with increasing nwnber trees diversity varied by site for the spring censuses of sampled in a stand. Data are from six of 1993-1995 usingANOVAon the Shannon stands in site 6 collected in September 1992. Weiner Diversity Index (Poole 1974).
Finally, we caculated a power analysis (Zar 1984) to determine the minimum detectable both white and black oak (fig. 2). Finally, difference for each census in insect density number of new species encountered with given the observed variation. The mean square increasing number of trees sampled per site error for block by treatment term from the appears to stabilize at about 30 to 35 trees for AN OVA for each separate census was used, with each host plant species (fig. 3). a= 0.05 and power (1-13) = 0.95. Together, these results suggested that a RESULTS sample of five trees per species per stand and six stands per site would be adequate to Preliminary Sampling characterize the variation in insect numbers and species composition within a site. Be Estimates of the mean number of insects per cause there was some evidence of differences tree per square meter of leaf area and standard among stands located on different slope posi deviation of that estimate appear to stabilize at tions, three stands were located on north- and approximately five to six trees at all six stands east-facing slopes (ELT 18) and three were sampled for both white (fig. 1) and black oak located on south- and west-facing slopes (ELT (not shown). For this particular sample, differ 17). ences in mean values between north- and south-facing slopes were not consistent for Sampling 1993-1995 either white or black oak (unpublished data). There were no significant main or interaction To determine the number of sampled stands effects involving treatment and block for necessary to characterize the variation in number of insects per leaf area on either white insects numbers at a site, the above estimates or black oak (table 1). ELT had a significant of the mean number of insects per tree for six impact (P = 0.00 1) on insect density on white trees at each of six stands were used to calcu oak and a significant interaction with year and late a mean value per stand. Means and stan census (both P = 0.001) (table 1). When differ dard deviation values for site 6 appear to stabi ences were significant, there were more insects lize after three to four stands were sampled for on trees on north- and east-facing slopes (ELT 335 ------
~ ~M@W~W------White Oak 5
C'CI Cll :;; 4
C'CI Mean 60 -Cll 3 --o-
C'\1 • Standard deviation 50 E "'~ 'c::i II) 2 ~ - Q. (J 40 -Cll "' II) c 1 -"~ .5"' 30 Whiteoak .... --o- 0 0 Black oak ~ 20 • 0 1 2 3 4 5 6 ,Q"' s z= 10 4 Black Oak
0,_--.---r-~r-~--~---r--~ 0 10 20 30 40 50 60 70 3 Number of trees sampled
--o- Mean Figure 3.- The relationship between number of 2 Standard deviation trees sampled and number of species of leaf chewing insects encountered in site 6 for both white and black oak. 1 • • Profile analysis for white oak (table 2) showed • ·-· that the effect of ELT increased from 1993 to 1994, and then again from 1994 to 1995, 0~------.---.----.---.--~ irrespective of season (i.e., the interaction effect 0 1 2 3 4 5 6 between ELT and year was significant for both Number of stands sampled per site contrasts) (fig. 4). The effect of ELTon insects feeding on white oak was also greater later in the season across all years: its effect increased Figure 2.- The relationship between the estimate from June to July (significant ELT effect), and of number of insects per tree per stand and then again from July to August censuses, but its standard deviation and increasing nwn not from May to June (table 2, fig. 4). ber of stands sampled in site 6 (September 1992) for both white and black oak. Insect densities, summed over all sites and all censuses, varied significantly across years (table 1), but more so for white than black oak 18) than on trees on south- and west-facing (fig. 5). For white oak, total numbers of insects slopes (ELT 17) (fig. 4). Stand as a main effect found on trees in 1994 and 1995 were twice was a marginally significant predictor of insect that in 1993; for black oak, totals in 1995 were abundance in white oak (P = 0.11) but not in 15 percent and 22 percent less than in 1993 black oak (P = 0.31). There was, however, a and 1994, respectively (fig. 5). Seasonal pattern significant interaction between stand and also varied by year. Seasonal variation in white census for both white and black oak P = 0.012 oak showed two pattems: either high abun and 0.013, respectively), demonstrating a stand dance in the spring and fall (1993) or an in effect for some censuses in both species, and a crease in abundance in spring to fall ( 1994 and significant year by stand interaction (P =0.014) 1995) (fig. 6). In contrast, seasonal pattem of in black oak (table 1). There were significant insect abundance on black oak was consistent interactions of block with census for white oak across years; abundance was highest in the (P = 0.052) and year for black oak (P = 0.069). spring (and 50 to 100 percent greater than white oak spring abundances), lowest in the midsummer, then increased again in the fall 336 (fig. 6).
337 337
1), 1),
F
oak oak
tree. tree.
four four
oak. oak.
in in
oak oak
the the
insect insect
and and
per per
that that
ELT ELT
by by
insect insect
and and
(table (table
black black
for for
stand. stand.
black black
significant significant
area area
black black
ELT ELT
for for
a a
by by
treatment, treatment, within
appears appears
on on
leaf leaf
block, block,
scale scale
overall overall
and and
calculating calculating
numerator numerator
revealed revealed
0.313 0.313
0.471 0.471
0.814 0.814
0.176 0.176
0.396 0.396
0.817 0.817
0.613 0.613
0.074 0.074
0.811 0.811
0.064 0.064
0.279 0.279
0.013 0.013
0.262 0.262
0.315 0.315
0.867 0.867
0.527 0.527
0.014 0.014
0.069 0.069
0.190 0.190
0.001 0.001
0.001 0.001
0.250 0.250
0.002 0.002
p p
p p
and and
not not
in in
analysis analysis
per per
by by
significant significant
nested nested
on on
the the
effect effect
a a
is is
in in
but but
white white
used used
the the
0.08) 0.08)
significant significant
velutina velutina
1.98 1.98
1.17 1.17
1.28 1.28
1.36 1.36
1.40 1.40
1.51 1.51
1.45 1.45
1.82 1.82 0.61 0.61
0.62 0.62
0.75 0.75
0.39 0.39
0.80 0.80
0.145 0.145
3.34 3.34
6.62 6.62
insects insects
interaction interaction
1.11 1.11
F F
1.18 1.18
0.77 0.77
0.06 0.06
F F
2.76 2.76
effect effect
37.17 37.17
analysis analysis
= =
significantly significantly
oak oak
Stand Stand
157.1 157.1
reported. reported.
of of
geographic geographic
but but
previous previous
(P (P
variation variation
velutina velutina
suggest suggest
freedom freedom
ELT ELT
are are
of of
of of
between between
4.46 4.46
the the
Quercus Quercus
MS MS
61.1 61.1
by by
79.4 79.4
white white
oak, oak,
effects. effects.
varied varied
second second
number number
399.7 399.7
552.4 552.4
235.8 235.8
denominator denominator
in in
Both Both
marginally marginally
results results
on on
on on
significant significant
site site
Quercus Quercus
the the
a a
As As
this this
degrees degrees
a a
1 1
2 2
4 4
2 2
2 2
is is
black black
12/60 12/60
6/29 6/29
significant significant
df df
102/801 102/801
34 34
6/66 6/66
differed differed
204/1602 204/1602
3/32 3/32
6/6 6/6
6/6 6/6
68/534 68/534
6/262 6/262
4/68 4/68
4/58 4/58
4/8 4/8
3/265 3/265
2/33 2/33
2/266 2/266
by by
Ndf/Ddf Ndf/Ddf
interaction interaction
of of
267 267
den den
species species
4). 4).
in in
was was
effects effects
had had
0.024). 0.024).
contribution contribution
MS MS
univariate univariate
= =
five five
subjects subjects within
subjects subjects
(P (P
effect effect
(table (table
census census
However, However,
census census
showed showed
abundance abundance
density density
numbers numbers there there
of of
ELT ELT
The The
subject subject
for for
are are
statistic. statistic.
p p
0.785 0.785
0.703 0.703
0.599 0.599
0.889 0.889
0.057 0.057
0.677 0.677
0.012 0.012
0.001 0.001
0.052 0.052
0.701 0.701
0.178 0.178
0.757 0.757
0.350 0.350
0.656 0.656
0.001 0.001
0.001 0.001
0.001 0.001
0.001 0.001
two two
pre-treatment pre-treatment
was was
MANOVA; MANOVA;
Ddf Ddf
test test
Within Within
0.110 0.110
p p
0.989 0.989
0.001 0.001
0.185 0.185
0.762 0.762
Between Between
years years
in in
F-statistic F-statistic
and and
entire entire
the the
ELT, ELT,
changes changes
not not
common common
Trace Trace
there there
alba alba
commu
black black
the the
1.16 1.16
1.34 1.34
possible possible
1.29 1.29
1.62 1.62
8.03 8.03
0.66 0.66
0.63 0.63
0.85 0.85
0.51 0.51
0.66 0.66
4.20 4.20
0.64 0.64
0.88 0.88 s s
9.14 9.14
0.63 0.63
at at
Ndf Ndf
commu
alba alba
F F
by by
values values
86.1 86.1
68.7 68.7
F F
abundance abundance
the the
1.30 1.30
significant significant
0.01 0.01
0.29 0.29
2.68 2.68
across across
133.7 133.7
14.42 14.42
that that
the the
For For
significantly significantly
were were
of of
most most
on on
Pillai Pillai
at at
testing testing
Quercus Quercus
entire entire
effects. effects.
1.67 1.67
five five
the the
example, example,
Quercus Quercus
MS MS
49.92 49.92
calculate calculate
patterns patterns
144.4 144.4
110.9 110.9
calculating calculating
169.4 169.4
453.4 453.4
showed showed
those those
OVA OVA
effect effect
varied varied
stand. stand.
on on
the the
to to
showed showed
2,081 2,081
those those
for for
the the
for for
AN AN
1 1
2 2
4 4
2 2
2 2
by by
significantly significantly
df df
of of
subjects subjects
42 42
significantly significantly
with with
ELT ELT
some some
126/894 126/894
12/76 12/76
6/82 6/82
6/37 6/37
252/1788 252/1788
3/40 3/40
616 616
616 616
84/596 84/596
6/293 6/293
Like Like
4/8 4/8 298 298
4/8 4/8
3/296 3/296
4/84 4/84
2/41 2/41
2/297 2/297
Ndf/Ddf Ndf/Ddf
based based
abundances abundances
oak, oak,
species species
species species
freedom freedom
census census
are are
3). 3).
and and
of of
varied varied
measures measures
between between
varied varied
error) error)
two two
white white
respectively, respectively,
MSE MSE
MSStand MSStand
MSStand MSStand
MSTxB MSTxB
MSTxB MSTxB
MSden MSden
three three
May May
species species
the the
contradicted contradicted
However, However,
abundance abundance
ELT. ELT.
(table (table
consistent consistent
plot plot
abundance abundance
For For
degrees degrees
the the
for for
and and
in in
species species
1 1
1 1
of of
and and
Repeated Repeated
species species
of of
level level
(C) (C)
species, species,
(whole (whole
all all
(B) (B)
one one
1.-
(subplot) (subplot)
level. level.
(T) (T)
0.001). 0.001).
(E) (E)
Stand Stand
B B
block, block, denominator, denominator,
statistic statistic
F-approximations F-approximations
block, block,
individual individual
= =
one one
"pre-treatment" "pre-treatment"
Insu:fficient Insu:fficient
(P (P
oak, oak,
of of
species species
1
nity nity
species species a a
necessarily necessarily
by by
community community
in in
nity, nity,
variation variation
YXCXStand YXCXStand
C C X
CXTXE CXTXE
YXCXTXE YXCXTXE
CXE CXE
YXCXE YXCXE
CXB CXB
YXCXB
CXT CXT
YXCXT
Analysis Analysis
Census Census Stand Stand
YXTXE YXTXE
YXStand YXStand
YXE YXE
YXB YXB
YXT YXT
YXC YXC Source Source
Year(Y) Year(Y)
ELT ELT
Source Source
TXE TXE
T T X
Treat Treat
Error Error
Block Block Table Table ~ ~~~©W~~------
D ELT17 E] ELT18
20 May 20 June
~ 15 ~ 15 ""~ ~ ~ ~ N- 10 10
rll -.....= ~ ~ rll 5 5 .....=
1993 1994 1995 1993 1994 1995
20 July 20 August
~ ~ ""~ 15 15 ~ ~ -~ N 10 10 rll .....= -~ ~ rll ..... = 5 5
1993 1994 1995 1993 1994 1995 Year Year
Figure 4.-E.ffect of ELT, year, and census on the mean(± SE) number of insects per leaf area on white oak.
338 Table 2.-Profile analysis for Quercus alba for the effect of either year or census (''Time") and ELT X Time White oak interaction ("ELT X Time") over the time periods 15 indicated on insect density per tree. 1993 Source F p D E'LI 1994 Contrast variable: Year 1- Year 2 =~ 1995 Time 171.6 0.001 = 10 ~- • =~ ELTXTime 18.4 0.001 - 1 Contrast variable: Year 2 -Year 3 ...(I) Time 2.4 0.123 -~ 5 ~ (I) ELTXTime 5.4 0.026 ~= Contrast variable: May -June Time 14.9 0.001 0 ELTXTime 1.0 0.323 May June July August
Contrast variable: June - July Month Time 5.0 0.026
ELTXTime 3.4 0.073
Contrast variable: July -August 15 Black oak Time 180.5 0.001
ELTXTime 4.5 0.040
=~ = 10 D 1993 ~- Ed 1994 =~ - 1995 M 5 • ...(I) -~ 40 ~ 5 (I) B B Quercus alba ~= 30 Quercus velutina
May June July August ~ g 10 Month ..5"'
0 Figure 6.-Seasonal variation by year in the 1993 1994 1995 mean (± SE) number of insects per leaf area Year in white oak and black oak.
Figure 5.-Annual variation in the mean (± SE) number of insects per leaf area on white and black oak. 339 ------~
+ 00 i!lllFJEIP Table 3 .-Repeated measures AN OVA testing possible pre-treatment effects on numbers of the five most common insect species per leaf area per tree. Analysis is the same as reported in table 1. NS. = non-significant at P>O.l 0.
Quercus alba Lithophane Chionodes Telphusa Spargo nothis Dichomeris Factor antennata sp. latifasciella pettitana ligulella
ELT (E) N.S. N.S. N.S. N.S. 0.005 Treatment (T) N.S. N.S. N.S. N.S. 0.017 Block (B) 0.080 N.S. N.S. N.S. 0.019 Stand (S) N.S. 0.061 0.023 N.S. 0.003 TXE N.S. N.S. N.S. N.S. N.S. Year(Y) 0.001 0.001 0.001 0.001 0.001 YXE N.S. N.S. N.S. N.S. 0.020 YXT N.S. N.S. N.S. N.S. N.S. YXB N.S. N.S. N.S. N.S. 0.003 YXS 0.003 0.006 N.S. N.S. N.S. YXTXE N.S. N.S. 0.031 N.S. N.S.
Quercus velutina Chionodes Telphusa unidentified Spargonothis Dichomeris Factor sp. latifasciella Tortricidae pettitana ligulella
ELT (E) 0.068 N.S. N.S. N.S. N.S. Treatment (T) N.S. N.S. N.S. N.S. N.S. Block (B) N.S. N.S. 0.078 N.S. 0.054 Stand (S) 0.035 0.006 0.016 N.S. 0.013 TXE N.S. N.S. N.S. N.S. N.S. Year(Y) 0.001 0.001 0.001 0.001 0.001 YXE N.S. N.S. N.S. N.S. N.S. YXT N.S. N.S. N.S. N.S. N.S. YXB N.S. 0.040 N.S. N.S. 0.066 YXS 0.002 0.004 N.S. N.S. 0.009 YXTXE N.S. N.S. N.S. N.S. N.S.
only some sites for the August-September values corresponds to the order in which sites census. As in white oak, when differences were sampled (sites are listed in sampling occurred, there were more insects on black oak order). The significant site by census interac trees of north- and east-facing slopes (ELT 18) tion in white oak is due to a similar pattem: a than on trees of south- and west-facing slopes range in site values occurs only in May, but (ELT 1 7) (25 percent more averaged across all again corresponds to the order in which sites sites in August censuses; not shown). were sampled (fig. 8). May is when the insect species composition changes most rapidly, and Site had a significant main effect for black oak total insect numbers decline over a vecy short (P = 0.005) and a significant site by census time period as leaves become fully expanded interaction for white oak P = 0.002). The site (RJM and JL, unpublished data). However, effect in black oak is due in part to consistently diversity (Shannon-Weiner index) did not de higher and lower insect numbers in site 6 and cline as sites were sampled in May for either site 2, respectively (fig. 7). The other contribut black or white oak (P > 0.15 for both species ing factor to the site effect is the great range in and all years). Site differences in insect abun insect numbers across sites for black oak dance in May on both black and white oak during the May censuses of 1993 and 1995 (fig. correspond to the time of acquisition of the sites 7, significant year by census by site interaction, by the MDC (fig. 9), with those sites under pattems by year not shown). The decline in control of the MDC longer having greater insect
340
by by
as as
the the
F-
341 341
with with
the the
black black
effect effect
to to
of of
effect effect
oak, oak,
varied varied
The The
differ
in in
the the
both both
tree. tree.
the the
and and
block block
of of
shows shows
misinter
one one
ELT. ELT.
block block
on on
existed existed
per per
of of
likely likely
block block
white white
oak). oak).
Trees Trees
of of be be
However, However,
and and
freedom freedom
some some
on on
the the
area area
(white (white
variable variable
of of
treatment treatment
site site
more more
model. model.
effect effect
0.720 0.720
0.110 0.110
0.436 0.436
0.018 0.018
0.253 0.253
0.081 0.081
p p
0.024 0.024
0.168 0.168
0.532 0.532
0.683 0.683
0.012 0.012
0.012 0.012
0.001 0.001
0.001 0.001
0.001 0.001
liguleUa, liguleUa,
for for
of of
could could
(black (black
leaf leaf
(although (although
was was
pre-treatment pre-treatment
numbers numbers
the the
design. design.
depending depending
within within
blocking. blocking.
per per
degrees degrees
were were
main main
census census
lack lack
keeping keeping
year year
in in
no no
later later
1.14 1.14
1.02 1.02
1.10 1.10
1.58 1.58
1.82 1.82
1.52 1.52
1.91 1.91
0.86 0.86
of of
2.51 2.51
0.96 0.96
0.80 0.80
2.27 2.27 8.34 8.34
F F
p p
0.547 0.547
interactions interactions 0.779 0.779
0.005 0.005
0.333 0.333
difference difference
40.22 40.22
effect effect
by by
velutina velutina
172.7 172.7
effects effects for for
account account
insect insect
are are
The The
nested nested
and and
insects insects
that that
original original
Dichomeris Dichomeris
insects insects
is is
velutina velutina
oak). oak). in in
species species
of of
original original
and and
number number
1.10 1.10
0.88 0.88
0.08 0.08
3.70 3.70
does does
of of
stand stand
F F
oak) oak)
there there
the the
sites sites
designed) designed)
Quercus Quercus
significant significant
the the
are are
Stand Stand
support support
oak) oak)
treatment treatment
in in
significant significant
the the
black black
to to
number number
no no
treatment treatment
Quercus Quercus
is is
Ddf Ddf
species, species,
that that
block block
on on
168/1602 168/1602
16/56 16/56
16/56 16/56
(white (white 84/801 84/801
6/23 6/23
48/168 48/168
48/168 48/168
24/84 24/84 3/26 3/26
24/84 24/84
2/27 2/27
56/534 56/534
6/262 6/262
3/265 3/265
variation variation
on on
1 1
Ndf/Ddf Ndf/Ddf 2/266 2/266
8 8
8 8
df df
pre-
but but
as as
treatments. treatments.
number number
28 28
Individual Individual
among among
were were
and and
a a
267 267
(black (black
that that
not not
originally originally
found found
scale scale
the the
F-statistic. F-statistic.
Ndf Ndf
subjects subjects
oak). oak).
(as (as
stand, stand,
unrelated unrelated Spatial Spatial
in in
census census
year year fact fact proscribed proscribed there there
such such
suggests suggests
variation variation
common common
but but
ences ences
preted preted
actual actual
We We
subject subject
the the
0.960 0.960
0.060 0.060
statistic. statistic.
0.680 0.680
0.022 0.022
0.023 0.023
0.001 0.001
0.002 0.002
0.818 0.818
0.001 0.001
0.045 0.045
0.310 0.310
0.796 0.796
0.001 0.001
0.001 0.001
0.001 0.001
p p
we we
p p
= =
geographic geographic
Within Within
Between Between
0.666 0.666
0.001 0.001
0.151 0.151
0.221 0.221
per per
P P
that that
of of
(30-50 (30-50
test test
total total
the the
some some
the the
effect effect
calculating calculating
(table (table
treat
for for
alba alba
1.16 1.16
1.31 1.31
1.53 1.53
0.64 0.64
1.82 1.82
1.17 1.17
0.66 0.66
8.26 8.26
0.72 0.72
2.26 2.26 8.97 8.97
0.85 0.85
of of
in in
86.1 86.1
68.7 68.7
for for
the the
F F
effect effect
oak, oak,
Trace Trace
1.42 1.42
1.26 1.26
0.73 0.73
133.7 133.7
alba alba
F F
by by
insect insect
insects insects
14.8 14.8
larger larger
's 's
0.05 0.05
in in
variation, variation,
the the
7 7
others others
= =
conclude conclude
-0.85, -0.85,
of of
year: year:
Quercus Quercus
(5. (5.
P P
in in
total total
black black
for for
oak oak
to to
Pillai Pillai
treatment treatment
1 1
8 8
8 8
Quercus Quercus
much much
df df
36 36
at at
a a
in in
and and
298 298
and and
the the
respectively, respectively,
OVA OVA
application application
and and
be be
while while
differences differences
on on
black black
to to
AN AN
change change
power power
that that
to to
Analysis Analysis
of of
-0.78 -0.78
estimates estimates
108/894 108/894
6/31 6/31 16/72 16/72
48/216 48/216
216/1708 216/1708
16/72 16/72
3/34 3/34
48/216 48/216
24/108 24/108
24/108 24/108
6/293 6/293
72/596 72/596 2/35 2/35
Ndf/Ddf Ndf/Ddf 2/297 2/297 3/296 3/296
or or
= =
white white
census census
based based
increase increase
prior prior
(r (r
have have
state state
small small
DISCUSSION DISCUSSION significant significant
for for
by by
are are
measures measures
Power Power
denominator, denominator,
5) 5)
white white
to to
be be
) )
present present
significant, significant,
2
statistical statistical
evidence evidence
sixfold sixfold
and and
m
would would
insects insects
0.004 0.004
be be
varies varies
no no
our our
to to
(table (table
either either
of of
relatively relatively
would would
per per
on on
and and
C) C)
a a
for for
enough enough
occurred. occurred.
4.-Repeated 4.-Repeated
Power Power
would would
(E) (E)
found found
(S) (S)
numerator numerator
approximations approximations
) )
2
twofold twofold
number number
had had ment ment
cases cases
6). 6). insects insects
differences differences
density density m
a a
Based Based
We We
have have
0.012 0.012
respectively). respectively).
numbers numbers
YXCXSXE YXCXSXE
YXCXStand YXCXStand
CXStand CXStand
YXCXE YXCXE
CXE CXE
CXSXE CXSXE
YXCXS YXCXS
cxs cxs YXE YXE
YXSXE YXSXE
YXStand YXStand
Source Source
Census( Census(
YXS YXS
YXC YXC
Year(Y) Year(Y)
SXE SXE
Site Site
Stand Stand
Error Error
ELT ELT
Source Source Table Table
~M©W~W------
Figure Figure
342 342
~ ~
~ ~
-
-
......
~ ~
~ ~
e e
= =
[/.l [/.l
[/.l [/.l
Q,j Q,j
~ ~
Q,j Q,j
"' "' ~ ~
-
Q,j Q,j
~ ~
~ ~ -
......
~ ~
~ ~
= =
e e
[/.l [/.l
[/.l [/.l
Q,j Q,j
~ ~
~ ~
"' "'
Q,j Q,j
~ ~
Q,j Q,j
~ ~
census census
15 15
10 10
10 10
15 15
7 7
0 0
5 5
5 5
.-Effect .-Effect
years years across
6 6
6 5 4 3 2 1 7 8 9 9 8 7 1 2 3 4 5 6
5 5
of of
site site
4 4
on on
for for
July July
3 3
estimates estimates
May May
Site Site
black black
2 2
oak. oak.
1 1
in in
Sites Sites
the the
7 7
mean mean
8 8
are are
shown shown
9 9
(± (±
SE) SE)
number number
in in
10 10
15 15
10 10
15 15
5 5
0 0
5 5
the the
6 5 4 3 2 1 7 8 9 9 8 7 1 2 3 4 5 6
order order
6 6
of of
insects insects
5 5
they they
4 4
were were
August August
per per
June June
Site Site
3 3
leaf leaf
sampled. sampled.
2 1 1 2
area area
for for
7 7
each each
8 8 9 9
~ ~
Ill Ill Ill Ill
.. ..
~ ~
Ill Ill Ill Ill
.. ..
the the
the the
343 343
of of
8 8
9 9
7 7
8 8
• •
• •
• •
7 7
•9 •9
• •
of of
• •
is is
and and
Ill Ill
~ ~
0 0
Ill Ill ~ ~
.. ..
0 0
number. number.
year year
point point
~ ~
""" """
Ill Ill
.. ..
P=0.004 P=0.004
~ ~
r=-0.85 r=-0.85
""" """
Ill Ill
.. ..
censuses censuses
1 1
2 2
1 1
• •
• •
2 2
• •
site site • •
auspices auspices
r=-0.79 r=-0.79
P=0.01 P=0.01
Each Each
~ ~
0 0 """ """
.. ..
~ ~
0 0 """ """
.. ..
May May
1993-1995)for 1993-1995)for
the the
oak oak
5 5
oak oak
• •
• •
5 5
between between
e3 e3 • •
•4 •4
in in
Conservation Conservation
acquisition acquisition
acquisition acquisition
~ ~
.., .., Ill Ill
.. ..
Ill Ill ~ ~
.. ..
.., ..,
oak. oak.
3,4 3,4
of of
years years of of
under under
of of
Black Black
White White
~ ~
0 0
.., .., .. ..
insects insects
~ ~
0 0
.., ..,
.. ..
the the
corresponding corresponding
black black
site site
Year Year
of of
Year Year
a a
its its
relationship relationship
Ill Ill ~ ~
6 6
N N
.. ..
~ ~
N N
Ill Ill
.. ..
• •
6 6
over over
of of
and and • •
Department Department
with with
The The
~ ~
0 0
N N
~ ~
.. ..
0 0
N N
density density
white white
3 3
2 2
4 4
6 6
5 5
7 7
6 6
9 9
7 7
5 5
8 8
11 11
9.-
10 10
12 12
labeled labeled both both
(calculated (calculated
mean mean
acquisition acquisition
Missouri Missouri
~ ~
~ ~
[IJ [IJ
c. c. ~ ~ 1-< 1-<
= =
[IJ [IJ
......
"""" """"
8 8
~ ~
~ ~
= =
[IJ [IJ
~ ~
[IJ [IJ
c. c. 1-< 1-<
e e
......
"""" """"
~ ~
Figure Figure
is is
so so
on on
the the
in in
May May
Why Why
mean mean
by by
a a
in in
those those
was was
sites sites
insects insects
fact fact
in in
num
black black
for for
more more
leading leading
census census
as as
the the
sampling sampling
be be
data). data).
of of
oak oak
ELT ELT
a a
shown shown
in in
sites sites
to to
The The
time time
that that
protection protection
and and
area area
total total
effect effect
therefore therefore
significant significant
depending depending
between between
(ELT), (ELT),
not not
are are
both both
spring, spring,
the the
spring spring
fact fact
the the
leaf leaf
proceeded. proceeded.
(and (and
is is
the the
ELT ELT
number number
(and (and
across across
years) years)
differences differences
in in
For For
the the
the the
the the
and and
Sites Sites
per per
3 3
located. located.
community community
significant significant
appeared appeared
estimates estimates
census, census,
that that
The The
the the
in in
relation relation
unpublished unpublished
position position
for for
May May
site site
census census
(for (for
on on
in in
sampled. sampled.
(no (no
the the
species species
Site Site
oak. oak.
varied varied
were were
species species
auspices auspices
change change
May May
thus thus
oak. oak.
early early
despite despite
insects insects
per per
the the
site site
did did
of of
slope slope
microsite microsite
oak oak
oak oak
strong strong
sites sites
the the
were were
of of
the the
not not
they they
of of
as as
and and
phenology. phenology.
suggests suggests
a a
(MOFEP (MOFEP
greatest greatest
white white
years) years)
site, site,
differed differed
in in
loss loss
black black
is is
in in
is is than than
their their
did did
2 2
both both
on on
white white
they they of of
to to
insects insects
greatest greatest
type type
under under
on on
across across
Perhaps Perhaps
which which
for for
(for (for
of of
number number
than than
there there
effect effect
are are
in in
both both
due due
stand stand
Stands Stands
soil soil
insects insects
6 6 5 4 3 2 1 7 8 9
order order
censuses censuses
8.-Effect 8.-Effect
numbers) numbers)
SE) SE)
came came
of of
in in
diversity diversity
by by
effect effect
trees trees
interaction), interaction),
clear. clear.
site site
site site
(± (±
the the
censuses censuses
white white
8 8
6 6
4 4
2 2
differences differences
sites sites
species, species,
climate climate
to to
artifact artifact
numbers numbers the the
site site
spring spring
not not
insect insect
oak oak
that that
the the bers bers
Finally, Finally, consistent consistent site site
for for
depending depending
independent independent
vary vary
vary vary
Figure Figure
whole. whole.
~ ~
~ ~ e':S e':S
e':S e':S
1-< 1-<
[IJ [IJ
~ ~
~ ~
e e
= = e':S e':S
[IJ [IJ
......
-
"""" """" ~ ~
- ~ ~
~M©W~~------
~ ~
bers bers
Year Year in in the the
time, time,
Table Table from from
4. 4. ment ment
Based Based
censuses censuses
suggestions suggestions
3. 3.
2. 2. acquisition acquisition
1952 1952
1952 1952
1938 1938
1952 1952
1944 1944
1944 1944
1938 1938
1938 1938
1925 1925
1. 1.
344 344
site site
The The
May May is is
necessarily necessarily
year. year.
be be
Because Because
from from
and and
area area
continue continue
and and
canopy, canopy,
mend mend two two
relatively relatively
Present Present
sample sample Relatively Relatively
small" small"
statistical statistical
stand stand species species
occur occur
stands stands stratified stratified
observed observed
of of
of of
5 5
grazing grazing
we we
of of
available. available.
.-Relation .-Relation
maintained. maintained.
on on
2 2
the the
insects insects
censuses censuses
host host
(± (±
significant significant
temporal temporal
Conservation. Conservation.
and and
one one
also also
and and
at at
our our
continued continued
or or
mean mean
SE) SE)
treatment treatment
per per
size size
turnover turnover
Site Site
sample sample
we we
about about
any any 2 2
4 4
9 9
7 7
3 3
5 5
6 6
8 8
black black
1 1
number number
plants, plants,
on on
unusually unusually
species species
and and
sampling sampling
high. high.
stand stand
for for
do do
high high
results, results,
power power
years years
site) site)
on on
number number
to to
suggest suggest
of of
would would
between between
both both
not not
Increased Increased
time time
white white
Quercus Quercus
variation variation
fire) fire)
1993-1995. 1993-1995.
other other
black black
and and
future future
numbers numbers
effect effect
2.8 2.8
3.9 3.9
3.0 3.0
5.9 5.9
4.5 4.5
4.1 4.1
4.1 4.1
3.3 3.3
6.1 6.1
Taken Taken
effects. effects.
size size
should should
We We
is is
suggesting suggesting
sampling sampling
know know
to to
among among
cannot cannot
effects, effects,
of of
species. species.
and and
of of
of of
we we
of of
jeopardize jeopardize
± ±
± ±
± ±
± ±
± ±
± ±
± ±
±0.6 ±0.6
±0.5 ±0.5
unclear unclear
by by
white white
year year
The The
distinguish distinguish
now now
that that
stands stands
oak oak
high high
0.2 0.2
0.4 0.4
0.3 0.3
0.4 0.4
0.3 0.3 (number (number
0.3 0.3
0.2 0.2
insects insects
oak oak
the the
the the
alba alba
of of
black black
make make
sampling: sampling:
ELT ELT
together, together,
what what
sampling sampling
be be
is is
of of
ELT ELT
reason reason
year. year.
species. species.
have have
the the
sampling sampling
be be
Missouri Missouri
are are
of of
but but
in in
oaks oaks
different different
incorporation incorporation
per per
considered. considered.
oak oak
for for
Moreover, Moreover,
should should
(fig. (fig.
insects insects
per per
extrapolated extrapolated
site site
the the
that that
four four
suggests suggests
factors factors
unusually unusually
reduction reduction
Quercus Quercus
of of
the the
square square
sufficient sufficient
four four
averaged averaged
In In
dominate dominate
site) site)
why why
7). 7).
"relatively "relatively
following following
trees trees
we we
6 6
11.4 11.4
results results
censuses censuses
5.9 5.9
7.9 7.9
6.1 6.1 7.4 7.4
7.8 7.8
(more (more 6.3 6.3
8.1 8.1
8.7 8.7
addition, addition,
power power
across across
Depart
should should
continue. continue.
times times
for for
can can
At At
recom
meter meter
cause cause
should should
num
± ±
± ±
± ±
± ±
± ±
± ±
± ±
± ±
± ±
velutina velutina
spatial spatial
per per
of of
that that
0.5 0.5
0.5 0.5
0.8 0.8
0.7 0.7
0.6 0.6
0.7 0.7
0.5 0.5
0.7 0.7
this this
1.2 1.2
over over
the the
of of
low low
site site
that that
the the
leaf leaf
a a
the the
is is
May May
August August
June June
May May
June June
July July August August
June June
6. 6.
July July May May
5. 5.
July July August August
Table Table
Census Census
7. 7.
vidual vidual
AN AN
number number
responses responses
Percent Percent
nity nity
abundance abundance
insect insect forestry forestry
Melanoplus Melanoplus
numbers. numbers.
post-treatment post-treatment differences differences
understand understand
is is
mation mation
regimes, regimes,
square square
particular particular
Estimates Estimates
tion tion
species species able able through through
tree tree each each Because Because
impacts impacts
more more Diapheromerajemorata, Diapheromerajemorata,
dance dance cratic cratic
Individual Individual
Sabine Sabine
should should
(mean (mean
difference difference
Symmerista Symmerista
1993 1993
1995 1995
1994 1994
1994 1994
1994 1994
1993 1993
1995 1995
1995 1995
1993 1993
6.-Minimum 6.-Minimum
important important
OVA) OVA)
1995 1995
1994 1994
1993 1993
in in census
growth growth
patterns. patterns.
(based (based
might might
square square
use use
meter meter
patterns patterns
numbers numbers
into into
increase increase
species, species,
for for
stimulea, stimulea,
be be
on on
should should
management management
of of
of of
would would
population population
of of
estimates estimates
in in
Quercus Quercus
31.3 31.3
41.3 41.3
28.2 28.2
37.5 37.5
30.5 30.5
a a
made made
10.3 10.3
species species
13.9 13.9
12.5 12.5
to to
of of
insect insect
their their
our our
Consumption Consumption
alternative alternative
damage damage
seriously seriously
leaf leaf
5.7 5.7
7.8 7.8
the the
8.5 8.5
8.7 8.7
suggest suggest
will will
error error
on on
sp.). sp.).
than than
treatment treatment
albifrons, albifrons,
= =
the the
estimating estimating
to to
amount amount
the the
detectable detectable
in in
represent represent
0.05 0.05
area area
laboratory laboratory
Such Such
species species
we we
year-to-year year-to-year
continue, continue,
compared compared
number number
sampling sampling
know know
entail entail
actually actually
for for
population population
and and
to to
treatments. treatments.
Psilocorsis Psilocorsis
will will
species species
Ignoring Ignoring
alba alba
(103) (103)
(556) (556)
(97) (97)
(302) (302)
(337) (337) (314) (314)
(246) (246) (471) (471)
(137) (137)
(313) (313)
(278) (278)
(386) (386)
and and
for for
recommend recommend
given given
translate translate
of of
block block
estimates. estimates.
that that
irruptions irruptions
abundance abundance
individual individual
reduce reduce
estimates estimates
the the
of of
power power
which which
leaf leaf
on on
individual individual
Heterocampa Heterocampa
difference difference
of of
is is
forest forest
effects. effects.
are are
observed observed
treatment treatment
leaf leaf
the the
given given
by by
chewing chewing
Lochmaeus Lochmaeus
sampling sampling
overall overall
rates rates
exhibit exhibit
because because
every every
relative relative
feeding feeding
to to
damage damage
changes changes
treatment treatment
known known
observed observed
Quercus Quercus
21.9 21.9
26.8 26.8
49.0 49.0
45.6 45.6
biology biology
(I (I
11.8 11.8
18.4 18.4
14.2 14.2
quercicella, quercicella,
variation variation
area area
our our
previous previous
overall overall
5.7 5.7 7.3 7.3
9.1 9.1
7.6 7.6
7.2 7.2
managements managements
insect insect
in in
-J3) -J3)
In In
then then
(e.g., (e.g.,
(Zar (Zar
parentheses. parentheses.
conducting conducting
year. year.
Given Given
values values
species species
insects insects
of of
insect insect
ability ability
= =
of of
addition, addition,
idiosyn
species' species'
consump
economic economic
trials) trials)
insect insect
to to
effects effects
effects effects
will will 0.95. 0.95.
by by
indi
variation variation
term term
(534) (534)
(205) (205)
1984) 1984)
(369) (369)
(303) (303)
(212) (212)
(190) (190)
(347) (347)
velutina velutina
(211) (211)
(500) (500)
(380) (380)
(345) (345)
(315) (315)
in in
is is
abun
commu
gutivitta, gutivitta,
can can
manteo, manteo,
go go
that that
insect insect
that that
per per
infor
avail
in in
these these
be be
to to
in in
for for
be be
total total
the the
on on
of of
the the
of of
a a it it
a a
1-
345 345
eds. eds.
We We
the the
Bull. Bull.
on on
Gen. Gen.
semi
R., R.,
an an
moths moths
Kathy Kathy
re
defolia
Andy Andy
1997. 1997.
Randy Randy
help help
J.A., J.A.,
Depart
1993. 1993.
came came
upland upland
tax
the the
MA: MA: Schultz, Schultz,
Res. Res.
In: In:
Service, Service,
the the
MO. MO.
U.S. U.S.
Kirk Kirk
Lepi
Station: Station:
Ozark Ozark
for for
Damond Damond
in in
William William
to to
Greig, Greig,
rearing. rearing.
planning. planning.
D.C. D.C.
and and
Brookshire) Brookshire)
and and
Environmental Environmental
Agricultural Agricultural
D.D. D.D.
Stephen Stephen
identified identified
Conservation. Conservation.
MN: MN:
Jack Jack
McGrath, McGrath,
in in
regarding regarding
comments comments
Dahms, Dahms,
and and
1991. 1991.
Forest Forest
Louis, Louis,
of of
Ecosystem Ecosystem
future. future.
Wold Wold
support support
acorns. acorns.
Boston, Boston,
Thomas, Thomas,
landscape landscape
Missouri Missouri
guide guide
symposium: symposium:
Dey, Dey,
Jakse, Jakse,
for for
who who
Nancy Nancy
Notman, Notman,
change change
CITED CITED
Brian Brian
Wolf, Wolf,
St. St.
vigor. vigor.
p. p.
Saturniidae) Saturniidae) to to
Paul, Paul,
Eric Eric
Missouri Missouri
(particularly (particularly
John John
p. p.
ecology ecology
and and
Sork Sork
R.; R.;
Eric Eric
J.M. J.M.
lowland lowland
the the
orangestriped orangestriped
of of
Butler, Butler,
advice advice
Liz Liz
Shifley, Shifley,
Experiment Experiment
field field
Missouri Missouri
40 40
caterpillar caterpillar
N.M.; N.M.;
Forest Forest
St. St.
and and
of of
the the
and and
3-5; 3-5;
P.B.; P.B.; Lill, Lill,
Evan Evan
A A
L.; L.;
496 496
Project Project
staff staff
and and
and and
infesting infesting
1318-1324. 1318-1324.
of of
America. America.
Passoa, Passoa,
Financial Financial
and and
discussions discussions
MO: MO:
reviewer reviewer
Linda Linda
thank thank
Agriculture, Agriculture,
Department Department
Forest Forest
Gonzalez, Gonzalez,
Biology, Biology,
Victoria Victoria
Jensen, Jensen,
present, present,
Ozark Ozark
approach approach
June June
22: 22:
to to
John John
Thomas, Thomas,
1984. 1984.
of of
Brian Brian
Catchpole, Catchpole,
Station. Station.
Collins, Collins,
growth growth
We We
indicators indicators
NC-193. NC-193.
Steve Steve
North North
Mifflin. Mifflin.
Robison, Robison,
Schultz, Schultz,
(Lepidoptera: (Lepidoptera:
late-season late-season
insects insects
LITERATURE LITERATURE
Jr. Jr.
support support
In: In:
Lill, Lill,
and and
B.L.; B.L.;
as as
ACKNOWLEDGMENTS ACKNOWLEDGMENTS
Whitney, Whitney,
1997 1997
grasslands grasslands
to to
past, past,
Hochwender, Hochwender,
lengthy lengthy
L.L.; L.L.;
1972. 1972.
oak oak
of of
thank thank
of of
Ecosystem Ecosystem
analysis. analysis.
Central Central
collection collection
Rep. Rep.
Floyd Floyd
draft. draft.
Missouri Missouri
Columbia, Columbia,
J. J.
for for
on on
Proceedings Proceedings
Mothershead, Mothershead,
Terry Terry
Conservation Conservation
Missouri Missouri
M.A.; M.A.;
we we
Katherine Katherine
C.V., C.V.,
anonymous anonymous
Cris Cris
Rick Rick
eastern eastern
Diane Diane
of of
the the
data data
an an
doptera doptera
Europe. Europe. natural natural
of of
Houghton Houghton
oakworm oakworm
Entomology. Entomology.
tion tion
25. 25.
Impact Impact
North North
search; search;
726. 726.
Forest Forest
experimental experimental
Department Department
Brookshire, Brookshire,
eds. eds.
Project: Project:
Experiment Experiment
Tech. Tech.
onomy onomy
grateful grateful
The The
thank thank
logistical logistical
Covell, Covell, Eberhardt, Eberhardt,
Coffelt, Coffelt,
Brookshire, Brookshire,
Brezner, Brezner,
Finally, Finally,
statistical statistical Connett Connett
for for
from from
caterpillars, caterpillars,
ment ment previous previous
are are
Kyllo, Kyllo,
and and
Jenson, Jenson,
Kristine Kristine
Hayes, Hayes, Stowe, Stowe,
Farrell, Farrell,
with with
We We
of of
the the
the the
10, 10,
tree tree
Li Li
may may
by by
might might
an an
g•owth g•owth
path path
each each
of of
figure figure
num
can can
on on
and and
rela
growth growth
are are
the the
will will
to to
tree tree
(see (see
in in
affect affect
Path Path
that that
effects. effects.
for for
under under
challenge challenge
T the the sampling sampling only only effects effects that that we we of of figure figure all all damage. damage. such such factors factors abundance abundance we we the the growth. growth. these these tree tree taxa taxa populations) populations) in in effects effects require require interaction. interaction. integration integration the the insects insects 1988) 1988) that that on on for for leaf leaf direct direct affect affect tree tree drawn drawn from from factors factors of of turn, turn, biggest biggest diagram diagram will will bird bird so so projects projects insect insect indirect indirect herbivores herbivores this this independent, independent, indirectly indirectly on on be be example example the the Thus, Thus, In In into into treatments treatments diversity diversity studied studied ~ ~ processes processes future future thoroughly thoroughly indirect indirect Our Our linking linking arise arise coefficients. coefficients. coordinated coordinated as as insects insects an an path path Horvitz Horvitz and and own own the the determining determining for for characteristics, characteristics, effects effects a a insect insect These These efforts efforts effect effect among among could could As As modify modify of of many many and and in in coefficients coefficients estimates estimates treatments treatments of of path path and and genetic genetic (including (including might might growth. growth. their their soil soil moderating moderating diversity diversity our our plea plea process. process. """""""'""" """""""'""" censuses censuses one one effects. effects. diagram diagram herbivorous herbivorous \ \ the the a a directly directly the the that that the the direct direct each each underlying underlying variables variables additional additional tree tree tree tree present present indirect indirect and and that that a a understood understood of of rsects~ rsects~ and and T7J..'"' T7J..'"' have have this this analysis analysis birds birds past past herbivorous herbivorous genetic genetic above above impact impact f f both both indirectly indirectly the the how how on on the the planning planning auspices. auspices. we we be be and and translate translate relationships. relationships. make make of of likely likely important important relates relates will will Schemske Schemske under under to to like like Tree Tree integrate integrate populations. populations. have have and and the the these these l.~ny l.~ny to to among among from from estimate estimate taxon taxon and and as as we we 10.-Path 10.-Path can can treatment treatment listed listed to to to to growth, growth, populations, populations, effect effect to to and and to to be be highly highly be be inMOFEP. inMOFEP. impact impact likely likely Insectivorous Insectivorous MOFEP MOFEP which which used used estimation estimation bers bers insect insect tree tree is is VogetaUo• VogetaUo• Figure Figure lead lead diagrams diagrams 10, 10, studied studied understand understand cooperative cooperative underlying underlying understand understand diagrams diagrams 1975 1975 Knowing Knowing will will tionship tionship considering considering may may on on want want for for factors factors However, However, direct direct shown shown processes processes insect insect growth growth pathogens, pathogens, to to directly directly sampling sampling the the It It Finally, Finally, understanding understanding integration, integration, ~@W~W------ The conservation of insects and their habi St. Paul, MN: U.S. Department of Agricul tats. 15th symposium of the Royal Entomo ture, Forest Service, North Central Forest logical Society of London. London: Academic Experiment Station: 26-40. Press: 213-236. Sparrow, H.R.; Sisk, T.D.; Ehrlich, P.R.; Hill, C.J. 1995. Linear strips of rain forest Murphy, D.D. 1994. Techniques and guide vegetation as potential dispersal corridors lines for monitoring Neotropical butterflies. for rain forest insects. Conservation Biology. Conservation Biology. 8: 800-809. 9: 1559-1566. Thomas, C.D.; Mallorie, H.C. 1985. Rarity, Legge, J.T.; Roush, R; DeSalle, R.; Vogler, A.P.; species richness and conservation: butter May, B. 1996. Genetic criteria for establish flies of the Atlas Mountains in Morocco. ing evolutionarily significant units in Biological Conservation. 33: 95-117. Cryan's Buckmoth. Conservation Biology. 10: 85-98. Tietz, H.M. 1972. An index to the described life histories, early stages and hosts of the Li, C. C. 1975. Path analysis. A Primer. Pacific Macrolepidoptera of the continental United Grove, CA: Boxwood Press. 210 p. States and Canada. Sarasota, FL: Allyn Museum of Entomology. 2 volumes. Littell, R.C.; Freund, R.J.; Spector, P.C. 1991. SAS system for linear models. Cary, NC: von Ende, C.N. 1993. Repeated-measures SAS Institute Inc. analysis: growth and other time-dependent measures. In: Scheiner, S.M.; Gurevitch, J., Marquis, R.J.; Whelan, C.J. 1994. Insectivorous eds. Design and analysis of ecological birds increase growth of white oak through experiments. New York, NY: Chapman and consumption of leaf-chewing insects. Ecol Hall: 113-137. ogy. 75: 2007-2014. White, J. 1995. Urbanization crowds out oaks. Parker, J.; Patton, R.L. 1975. Effects of drought California Agriculture. 49: 5-6. and defoliation on some metabolites in roots of black oak seedlings. Canadian Journal of Zar, J.H. 1984. Biostatistical analysis. 2d ed. Forest Research. 5: 457-463. Englewood Cliffs, NJ: Prentice-Hall. 718 p. Poole, R.W. 1974. An introduction to quantita tive ecology. New York, NY: McGraw-Hill. 532p. Rexrode, C.O. 1971. Insect damage to oaks. In: The oak symposium; 1971 August 16-20; Margan town, WV. U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 129-134. Schemske, D.W.; Horvitz, C.C. 1988. Plant animal interactions and fruit production in a neotropical herb: a path analysis. Ecology. 69: 1129-1137. Sheriff, Steven L.; He, Zhuogiong. 1997. The experimental design of the Missouri Ozark Forest Ecosystem Project. In: Brookshire, Brian L.; Shifley, Stephen R., eds. Proceed ings of the Missouri Ozark Forest Ecosystem Project symposium: an experimental ap proach to landscape research; 1997 June 3- 5; St. Louis, MO. Gen. Tech. Rep. NC-193. 346