Forest Ecology and Management 201 (2004) 327334

www.elsevier.com/locate/foreco

Comparison of methods for sampling Thysanoptera on basswood

(Tilia americana L.) trees in mixed northern hardwood
deciduous forests
S.M. Wernera,*, E.V. Nordheimb, K.F. Raffaa
a

Department of Entomology, 345 Russell Laboratories, University of Wisconsin, Madison, WI 53706, USA
Department of Statistics and Forest Ecology and Management, University of Wisconsin, Madison, WI 53706, USA
Received 10 May 2004; received in revised form 13 July 2004; accepted 15 July 2004

Abstract
Canopy arthropods play integral roles in the functioning, biodiversity, and productivity of forest ecosystems. Yet quantitative
sampling of arboreal arthropods poses formidable challenges. We evaluated three methods of sampling the introduced basswood
thrips, Thrips calcaratus Uzel (Thysanoptera: Thripidae), from the foliage of basswood canopies with respect to statistical
variability and practical considerations (legal, economic and logistical accessibility). All three methods involved removal of
foliage, which was performed using a pole-pruner, shotgun, and certified tree-climber. We also tested a fourth method, in which
the tree-climber enclosed samples in a plastic bag to estimate losses that occur when branches fall to the ground, even though this
is often not practical. The climber plus bag and pole-pruning methods obtained the highest numbers of thrips. Mean number of
larval thrips did not vary significantly among the three main sampling methods. Site had a stronger effect on the number of larval
thrips obtained than on the number of adults. A significant method by site interaction was observed with adults but not larvae.
Significant collection date (which corresponds to thrips life stage) by site interaction was also observed. We regressed sampling
methods to determine if the number of thrips obtained using one method can be used to predict the number obtained with another.
Tree-climber and pole-pruner data were highly predictive of each other, but shotgun data cannot be used to estimate other
methods. Pole-pruning is the most cost-effective and legally permissible technique, but is limited to trees with accessible lower
branches. The shotgun method is cost-effective and useful in sampling trees at least up to 27 m, but is prohibited close to human
activity. The tree-climber is effective and broadly applicable, but incurs the highest costs. This study shows the need to evaluate a
variety of techniques when sampling arboreal insects with respect to predictability, pragmatics and life stages.
# 2004 Elsevier B.V. All rights reserved.
Keywords: Thysanoptera; Sampling methods; Canopy; Tilia americana

* Corresponding author. Present address: DCNR, Forest Pest

Management, 208 Airport Drive, 2nd Floor, Middletown,
PA 17057, USA. Tel.: +1 717 651 9113.
E-mail address: shahlawerner@yahoo.com (S.M. Werner).

1. Introduction
An understanding of the abundance and diversity
of canopy arthropods can provide insight into basic

0378-1127/$ see front matter # 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2004.07.014

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S.M. Werner et al. / Forest Ecology and Management 201 (2004) 327334

ecosystem processes and biodiversity (Hammond and

Miller, 1998; Richardson et al., 1999). Population data
may also be used to monitor the impacts of introduced
species (Orwig and Foster, 1998), herbivory (Batzer et
al., 1995; Jackson, 1996), forest management practices
(Schowalter, 1995; Watt et al., 1997; Su and Woods,
2001), and natural or anthropogenic disturbance (Schowalter, 1994). Unfortunately, quantitative sampling of
arboreal arthropods poses a number of logistical and
statistical challenges.
A number of indirect methods have been attempted,
such as sampling frass, alternate life stages, and plant
injury (Allen et al., 1986; Parker and Skinner, 1993;
Zandt, 1994; Liebhold et al., 1995; Liang et al., 1998).
However, these methods do not always accurately
estimate the targeted life stage, provide only relative
estimates, and are not readily amenable to repeated nondestructive sampling (Zandt, 1994; Liang et al., 1998).
A variety of direct methods have also been employed,
each with certain advantages and disadvantages. Tree
felling is commonly used (Shepherd and Brown, 1971;
Batzer et al., 1995), but this is destructive, costly, and
eliminates the possibility for repeated sampling. Sampling from scaffolding and gondolas is non-destructive,
amenable to repeated sampling, and allows direct access
for detailed in situ observations. However, these methods
are costly (Jackson, 1996), and restricted to certain types
of canopies and terrain (Lowman, 2001; Stork, 2001).
They likewise restrict the number of tree- or site-level
samples, which reduces their applicability to studies of
population dynamics and habitat associations. Sampling
with a pole-pruner can generate large sample sizes, but
also has limitations. Since insects are often clustered in
distribution (Kuno, 1991; Pena and Duncan, 1992),
methods that only sample the lower portion of the canopy
may introduce bias. Sweney and Jones, 1975 sampled
branches with a rifle, but the extent to which this method
affects absolute arthropod numbers is unknown. Herms
et al., 1990 found that a D-vac more effectively sampled
adult Hemiptera, but a beating tray was optimal for early
instars. Branch disturbance created by beating, D-vac
suction, or shooting results in some error. Despite the
need to understand issues associated with arboreal insect
sampling, relatively few studies have addressed this
subject directly, particularly from the perspective of
comparison of sampling methods.
American basswood, Tilia americana L., is a major
component of northern hardwood forests throughout the

northeastern United States and southeastern Canada

(USDA FS, 1991). It often occurs in mixed stands,
and is associated with red maple, Acer rubrum L., sugar
maple, A. saccharum Marsh., northern red oak, Quercus
rubra L., eastern hemlock, Tsuga canadensis (L.) Carr.,
and yellow birch Betula alleghaniensis Britton. Mature
trees can grow up to 43 m tall in closed stands, and often
have long straight boles lacking lower branches (Burns
and Honkala, 1990; USDA FS, 1991). Basswood provides habitat and pollen for wildlife, helps maintain soil
quality (Burns and Honkala, 1990) and has several
economic uses (Burns and Honkala, 1990; USDA FS,
1991). Recently, forest monitoring in the Great Lakes
region has recorded a significant increase in basswood
crown dieback, mortality and defoliation (http://
www.na.fs.fed.us/spfo/fhm/index.htm).
Thysanoptera, commonly known as thrips, are an
important component of forest canopies, and some can
cause extensive damage to trees (Lewis, 1973; Parker et
al., 1988; Raffa and Hall, 1989; Kolb and Mccormick,
1993; Mound, 1999). At least five species, including
three herbivores: Thrips calcaratus Uzel (introduced
basswood thrips), Neohydatothrips tiliae (Hood) (native
basswood thrips), Taeniothrips inconsequens (Uzel)
(pear thrips), and two predators: Leptothrips mali (Fitch)
(black hunter thrips) and Aeolothrips melaleucus Haliday, are known to occur on American basswood (Stannard, 1968; Raffa and Hall, 1989; Raffa et al., 1992;
Rieske and Raffa, 1996). Only T. calcaratus has been
associated with defoliation and dieback in basswood
(Raffa and Hall, 1989).
T. calcaratus is univoltine and parthenogenic in
North America, comprised exclusively of diploid
females that reproduce asexually through thelytoky
(Ananthakrishnan, 1990). Adults emerge in early
spring, feed on developing buds, and oviposit on
the undersides of leaf midveins. Emerging larvae feed
on foliage for approximately two weeks before dropping to the ground to pupate within earthen cells
(Lewis, 1973; Raffa et al., 1992; Rieske and Raffa,
1996; Parker et al., 1992).
Our objective was to compare four sampling methods (pole-pruning, shotgun, tree-climbing, and treeclimbing plus bagging) in terms of absolute numbers,
statistical variability and economic, legal, and logistical accessibility. We also tested whether correction
factors among procedures are applicable when circumstances require multiple methods. This experi-

S.M. Werner et al. / Forest Ecology and Management 201 (2004) 327334

ment was conducted as part of a larger study on

impacts of the introduced basswood thrips on forest
health in the Great Lakes region of North America.

2. Materials and methods

The study area consisted of four northern hardwood
forest sites in northern Wisconsin (Table 1). Locations
ranged from 45.2 to 46.28 north latitude, and from 88.6 to
91.28 west longitude (Table 1). We sampled each site on
11 and 25 May 2001, which correspond to periods when
adults and larvae are most abundant, respectively (Raffa
et al., 1992). Tree diameter ranged from 17 to 43 cm at
1.5 m height (Table 1). A sample consisted of the outermost three buds or leaf clusters from a small branch.
Samples were placed in a plastic bag containing 70%
ethanol, and extracted in the laboratory using a suction
funnel containing a white 812 cup coffee filter. Thrips
were removed from the filter with a fine paintbrush and
examined under a compound microscope.
The following methods were compared for relative
numbers of thrips in a given foliage sample: (1) an
1.83 m pole-pruner with up to four additional 1.83 m
extensions; (2) a 20-gauge (15.90 mm bore diameter)
shotgun; (3) tree-climber, branch allowed to fall to the
ground and (4) tree-climber, branch immediately
enclosed in plastic bag. Our primary focus was to
compare the first three sampling methods. The last
method was included to estimate numbers of insects
lost due to branch disturbance using other methods,
but has limited utility because it is highly laborious
and introduces bias in the branches that may be
selected for sampling. Eight trees per method were
Table 1
Summary of study site locations and diameter of sampled basswood
trees in Wisconsin
Site

Location

FHMa 4508868
NAMPb #6
NAMP #16
NAMP #14

458440 N,
458180 N,
468280 N,
458510 N,

Diameter of
sampled trees (cm)
888560 W
888590 W
908210 W
918140 W

33.15,
23.50,
25.47,
20.21,

41.94
20.72
42.95
17.38

FHM refers to a plot that was included in the national USFS

Forest Health Monitoring network.
b
NAMP refers to a plot that was part of the North American
Maple Project, a 10-year study to investigate the causes of sugar
maple decline.

329

sampled during each collection, distributed equally

among the four sites.
A split plot analysis within PROC MIXED (SAS
Institute, 1999) was used to analyze the numbers of T.
calcaratus obtained. Fixed effects included site, collection date, and sampling method. Individual trees
(two trees/site) were included as a random effect and
viewed as the whole plot error term for testing the main
effect of site. The residual error was used for testing all
other factors, including interactions. Data were analyzed using either the three main sampling methods
(shotgun, tree-climber, and pole-pruner) or all four
methods. Because we observed a significant effect of
collection date, and significant interactions between
collection date with site and method (see Section 3),
we also analyzed the two time periods separately.
Because the first sampling period consisted almost
entirely of adults and the second consisted almost
entirely of larvae (see Section 3), we limited singledate analyses to the respective dominant life stages.
Variance increased with the mean, so a log10(x + 1)
transformation was applied to normalize the data.
We explored the relationships among methods and
the capability of predicting the numbers of thrips that
would have been obtained by one method from the
numbers obtained using another method. Specifically,
numbers of thrips obtained using one of the three main
sampling methods were regressed against each other.
All pairwise combinations were considered. Regressions were computed separately for adults and larvae.
A cost comparison of the methods was performed.
Costs were separated into fixed (independent of sampling intensity) and variable (intensity-dependent)
costs. Fixed costs included equipment, and variable
costs included labor, vehicle rental, and supplies. We
also compared the methods based on the costs required
to drive to distant study areas, as this is often required for
ecological studies. Because driving time is included in
the hourly charges of field assistants, and their fees vary
substantially with expertise required for each method,
we included one scenario involving no travel and one
with 200 km round-trip travel.

3. Results
We obtained 3047 thrips representing five species
(Table 2). Over 99% of these were T. calcaratus. Other

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S.M. Werner et al. / Forest Ecology and Management 201 (2004) 327334

Table 2
Total numbers and species of Thysanoptera obtained from basswood
canopy foliage
Species

5/10/01

5/25/01

Adults Larvae Adults Larvae

Herbivores
Suborder Terebrantia
Neohydatothrips tiliae
0
(Hood)
Taeniothrips inconsequens
0
(Uzel)a
Thrips calcaratus Uzela
547
Predators
Suborder Terebrantia
Aeolothrips melaleucus
Halidaya
Suborder Tubillifera
Leptothrips mali (Fitch)
a

12

105

2365

Indicates introduced species.

species obtained in low abundance were N. tiliae,

T. inconsequens, L. mali, and A. melaleucus. Based
on this distribution, we limited our statistical analysis
to T. calcaratus. 98% of the T. calcaratus obtained
during the first collection were adults, and 96%
obtained during the second collection were larvae
(Table 2). This pattern was observed for all sampling
methods and at all sites.
Samples obtained by a tree-climber and bagged
before falling to the ground caught an average of 71
thrips per sample, followed by the pole-pruner, treeclimber, and shotgun, which caught about 64, 51 and
40% as many thrips per sample, respectively (Table 3).
The shotgun and tree-climber methods caught significantly fewer thrips than the tree-climber with bag
method. When adults were considered separately,
the pole-pruner method obtained slightly more thrips
than the tree-climber with bag method, although this
difference was not significant (Table 3). The tree

climber with bag method caught significantly more

larvae than the other methods. All methods showed
relatively high standard errors.
When all four methods were considered across both
collection dates, the tree (within site) factor was not
significant. The 2Residual Log Likelihood value for
tree was significant when evaluated on a chi-squared
table. There were significant effects of collection date
(P = 0.0001) and sampling method (P = 0.0001).
A strong interaction between collection date and
site was also observed (P = 0.0003) (Table 4). The
site by method interaction was significant overall (P =
0.0420), and when adults were considered separately
(P = 0.0107). In addition, there was a significant
interaction for site by collection date by method (P
= 0.0056). This might be due to the large main effect
of method for larvae (P = 0.0017) and by the fact that
the site by method interaction was much more pronounced during the first (adult thrips) than second
(larvae) collection. Because of the strong effect of
collection date, we analyzed the data separately for
each insect life stage. There was a significant effect of
foliage sampling method on the number of adults (P =
0.0018) and larvae (P = 0.0017) when all four methods
were considered. The effect of site was significant for
larvae (P = 0.0284) but not for adults (Table 4).
The random effect of tree within site was not significant when the three primary methods were compared. The 2Residual Log Likelihood value for tree
was significant when evaluated on a chi-squared table.
The effects of collection date (P = 0.0001) and sampling method (P = 0.0079) significantly differed in
terms of the number of thrips obtained, when numbers
of adults and larvae were combined (Table 5). In
addition, there was a significant pairwise interaction
between collection date and site (P = 0.0001) and a
significant method by site interaction (P = 0.0246).
Due to the significant main effect of collection date

Table 3
Comparison of mean ( S.E.) T. calcaratus obtained from basswood canopy foliage using four sampling methods
Sampling method

Mean/samplea
(N = 16)

S.E.

Mean adults/sample
5/10/01 (N = 8)

S.E.

Mean larvae/sample
5/25/01 (N = 8)

S.E.

Shotgun
Pole pruner
Tree-climber
Climber + bag

28.75b
45.75ab
36.12b
71.38a

5.68
10.31
7.90
16.04

11.38a
22.13a
14.50a
20.38a

2.66
6.52
4.70
3.48

46.13
69.38
57.75
122.38

6.72
15.91
10.56
18.63

b
b
b
a

Mean thrips/sample within a column followed by the same letter are not significantly different (Tukeys L.S.D., 95% CI).

S.M. Werner et al. / Forest Ecology and Management 201 (2004) 327334

331

Table 4
Split plot analysis to evaluate fixed effects, considering four methods
of sampling thrips in basswood canopies

Table 5
Split plot analysis to evaluate fixed effects, considering three
methods of sampling thrips in basswood canopies

Source of variation

Source of variation

d.f.

2
3
1
6
2
3
6

6.24
4.55
178.39
3.14
1.47
15.17
0.34

0.0079
0.0886
0.0001
0.0246
0.2547
0.0001
0.9067

d.f.

Overall
Method
Site
Collection datea
Method  site
Method  collection date
Collection date  site
Method  site  collection date

3
3
1
9
3
3
9

17.20
4.53
298.43
2.33
2.59
8.68
3.46

0.0001
0.0894
0.0001
0.0420
0.0730
0.0003
0.0056

Overall
Method
Site
Collection datea
Method  site
Method  collection date
Collection date  site
Method  site  collection date

Adults (5/10/01)
Method
Site
Method  site

3
3
9

9.39
4.35
4.31

0.0018
0.0947
0.0107

Adults (5/10/01)
Method
Site
Method  site

2
3
6

11.88
4.95
4.17

0.0040
0.0782
0.0336

Larvae (5/25/01)
Method
Site
Method  site

3
3
9

9.51
9.27
1.81

0.0017
0.0284
0.1669

Larvae (5/25/01)
Method
Site
Method  site

2
3
6

0.85
10.40
0.93

0.4627
0.0233
0.5204

Collection date refers to adults collected on 5/10/01, and larvae

collected on 5/25/01.

Collection date refers to adults collected on 5/10/01, and larvae

collected on 5/25/01.

and interactions between collection date with site

and method, we analyzed data separately for adults
obtained on 10 May and larvae obtained on 25 May.
There was a significant site effect on the number of
larvae (P = 0.0233), but not on the number of adults
obtained. In contrast, there was a significant effect
of method on the number of adults (P = 0.0040),
but not larvae. There was a significant interaction
between method and site for adults (P = 0.0336),
but not larvae.
The regression analyses indicate that the numbers
of thrips obtained by pole-pruner and tree-climber

methods were highly correlated, but neither was correlated with shotgun data (Table 6). This pattern was
observed for both adults and larvae. The predictability
between methods was highest when pole-pruner data
were regressed on tree-climber data for adults (R2 =
0.73) and larvae (R2 = 71). Although the regressions of
shotgun data on other methods were higher for adults
than larvae, the results for the slope term were never
significant at P < 0.05, and thus the shotgun results are
not well-predicted by any of the other methods.
The economic analysis indicated that pole-pruning
was the most cost effective method of sampling

Table 6
Regression of methods used to sample thrips in basswood canopies
Methods

F (P > F)

R2

Slope (S.E.)

Intercept (S.E.)

Intercept T (P > T)

Adults (5/10/01)
Shotgun vs. pole-pruner
Shotgun vs. tree-climber
Pole-pruner vs. tree-climber

3.17 (0.13)
5.18 (0.06)
16.33 (0.01)

0.35
0.46
0.73

0.70 (0.39)
0.83 (0.37)
0.83 (0.21)

0.11 (0.51)
0.09 (0.41)
0.04 (0.27)

0.22 (0.84)
0.21 (0.84)
0.15 (0.88)

Larvae (5/25/01)
Shotgun vs. pole-pruner
Shotgun vs. tree-climber
Pole-pruner vs. tree-climber

0.87 (0.39)
1.52 (0.26)
14.58 (0.01)

0.13
0.20
0.71

0.19 (0.20)
0.35 (0.29)
0.56 (0.15)

1.31 (0.36)
1.03 (0.50)
0.74 (0.26)

3.69 (0.01)
2.07 (0.08)
2.84 (0.03)

Pairwise comparisons performed to determine predictabilities among methods and to evaluate applicability of applying a correction factor when
more than one sampling method is used.

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S.M. Werner et al. / Forest Ecology and Management 201 (2004) 327334

Table 7
Economic comparison of methods used to sample basswood canopies for thrips (USD, 2001)a
Method

Certified
Tree-climber
Shotgun

Pole-pruner w/two
1.8 m extensions
a

Fixed costs
(equipment) ($)

650
250

166

Variable costs
(labor, supplies, travel)

$20 h1 driving, set-up

$40 h1 climbing
$5/box ammunition (30 shells)
$8/day + $0.20/1.61 km vehicle rental
$1362/month graduate stipend (8.51 h1)
$8/day + $0.20/1.61 km vehicle rental
$1362/month graduate stipend (8.51 h1)

Avgerage time (h)

to collect 30
samples from 10 trees

Cost to collect 30 samples

from 10 trees ($)
200 km driving
(2.27 h)

No driving

105.40

60.00

74.32

41.48

69.32

27.83

Calculations based on fixed and variable costs for Wisconsin in 2001. Results will vary based on in-house resources, wages and inflation.

foliage, followed by the shotgun and tree climber

(Table 7). Tree-climber costs are relatively high due
to the equipment costs and higher hourly wages
incurred, especially if a certified tree-climber must
travel to distant sites. Thus, the absolute and relative
differences vary with proximity to the sites. Moreover,
these specific fixed and variable costs are likely to vary
among agencies, regions, and years, and so are of
primarily relative benefit.

4. Discussion
These results suggest that the optimal method for
sampling canopy insects can vary with the objectives
of a given study. For example, if the goal is to obtain
the maximum number of insects, the tree-climber with
bag method is most effective, as it reduces losses due
to branch disturbance (Table 3). However, if it is
necessary to sample various sections of the canopy,
and the objective is to compare numbers of thrips
among trees or sites, the shotgun or tree-climber (unbagged sample) methods might be preferable because
of the larger number of samples that can be obtained
(i.e., pole-pruners often can only reach the lower
sections of the canopy). The final decision would
likely incorporate factors such as proximity to human
activity, research personnel, and budget (Table 7).
Another important consideration is the amount of time
required for each method relative to the phenologies of
the target species. If it is necessary to sample both
multiple trees per site and multiple sites, it may be

difficult to comprehensively sample species that have

brief seasonal activity periods by tree-climbing. Treeclimbing is much more time-consuming than other
methods, especially when care must be taken not to
damage the tree (e.g. climbing spikes may not be
permitted).
It may be necessary to sample foliar insects using
more than one method within the same study, due to
factors related to logistics, economics, and legal
access. For instance a pole-pruner is unable to reach
branches on trees that lack lower branches, and a
shotgun may not be permitted close to human activities. If a correction factor can be applied to methods
that vary in a statistically predictable way (i.e. no
significant interactions among methods and other
factors), using different methods can provide
increased sample sizes and statistical replication. This
study found the tree-climber and pole-pruner methods
could predict results between one another, but neither
method could predict shotgun samples (Table 6). This
might be because these two methods create less branch
disturbance than shotgun sampling.
In several cases results changed with insect development stage. For instance, site factors significantly
affected the number of larvae but not the number of
adults. This suggests that there may be lower variability in factors that strongly affect T. calcaratus
adults, such as soil type or tree architecture, than
factors that affect larvae, such as light exposure.
Method affected the number of adults but not larvae
obtained using the three primary methods. This may
be explained by differences in activity and distribution

S.M. Werner et al. / Forest Ecology and Management 201 (2004) 327334

between adult and larval thrips. Larvae are more

sessile and appear less selective about feeding sites
than adults, which emerge early in the spring and must
select the branches where buds have opened first.
These parts of the tree tend to be more exposed and
to extend further from the main portion crown, thus
making them more accessible to the pole-pruner and
tree-climber than the shotgun method (where branches
are usually selected from a thick clump in the middle
of the crown). The decreased mobility and increased
visibility of larvae may also increase their susceptibility to natural enemies on foliage. The significant
site by method interactions we observed for adults (but
not larvae) may likewise reflect differences in mobility
and the effects of canopy architecture on their ability
to respond and fall off when foliage is disturbed.
Caution must be applied in interpreting results of
canopy sampling, as all methods have biases and
limitations. A clear objective, e.g. obtaining a maximum number of insects vs. making comparisons
among sites, is needed for good experimental design.
Additionally, recording the precise position in the
canopy from which each sample was derived might
facilitate comparisons among sampling methods, as
insect numbers may vary among strata of the tree
canopy (Batzer et al., 1995).

Acknowledgements
We thank tree-climber Sean Gere, lab assistant
Prima Chambers, UW-Madison, and the Wisconsin
Department of Natural Resources for providing assistance in the field. We thank computer consultant Peter
Crump, UW-Madison, College of Agricultural and
Life Sciences for assisting with data analysis. We
thank Sueo Nakahara, USDA Agricultural Research
Service, Systematics Entomology Laboratory, Beltsville, MD, for providing taxonomic support. We thank
UW Madison professors Michael Adams, Department
of Botany, Richard Lindroth, Department of Entomology, Craig Lorimer, Department of Forest Ecology
and Management, Daniel Mahr, Department of Entomology and Daniel Young, Department of Entomology, for input on experimental design and manuscript
review. The USDA FS Forest Health Monitoring
program and the UW-Madison College of Agricultural
and Life Sciences provided support for this project.

333

The helpful comments of two anonymous reviewers

are greatly appreciated.
References
Allen, D.C., Abrahamson, L.P., Eggen, D.A., Lanier, G.N., Swier,
S.R., Kelley, R.S., Auger, M., 1986. Monitoring spruce budworm (Lepidoptera: Tortricidae) populations with pheromonebaited traps. Environ. Entomol. 15, 152165.
Ananthakrishnan, T.N., 1990. Reproductive Biology of Thrips,
Indira Publishing House, Oak Park, Michigan (p. 158).
Batzer, H.O., Martin, M.P., Mattson, W.J., Miller, W.M., 1995. The
forest tent caterpillar in aspen stands: distribution and density
estimation of four life stages in four vegetation strata. For. Sci.
41, 99121.
Burns, R.M., Honkala, B.H., 1990. Silvics of North American Trees,
vol. 2. Hardwoods, USDA FS Agricultural Handbook, 654 pp.
Hammond, P.C., Miller, J.C., 1998. Comparison of the biodiversity
of Lepidoptera within three forested ecosystems. Ann. Entomol.
Soc. Am. 91, 323328.
Herms, D.A., Nielsen, D.G., Sydnor, T.D., 1990. Comparison of two
methods for sampling arboreal insect populations. J. Econ.
Entomol. 83, 869874.
Jackson, R.V., 1996. A new technique for accessing tree canopies to
measure insect herbivory. Aust. J. Entomol. 35, 9395.
Kolb, T.E., Mccormick, L.H., 1993. Etiology of sugar maple decline
in 4 Pennsylvania stands. Can. J. For. Res. 23, 23952402.
Kuno, E., 1991. Sampling and analysis of insect populations. Annu.
Rev. Entomol. 36, 285304.
Lewis, T., 1973. Thrips: Their Biology, Ecology and Economic
Importance, Academic Press, New York.
Liang, Q., Otvos, I.S., Bradfield, G.E., 1998. Pupal sampling of the
western hemlock looper, Lambdina fiscellaria lugubrosa (Hulst)
(Lep., Geometridae) using burlap traps. J. Appl. Entomol. 122,
8588.
Liebhold, A.M., Elkinton, J.S., Zhou, G., Hohn, M.E., Rossi, R.E.,
Boettner, G.H., Boettner, C.W., Burnham, C., McManus, M.L.,
1995. Regional correlation of gypsy moth (Lepidoptera: Lymantriidae) defoliation with counts of egg masses, pupae, and male
moths. Environ. Entomol. 24, 193203.
Lowman, M.D., 2001. Plants in the forest canopy: some
reflections on current research and future direction. Plant Ecol.
153, 3950.
Mound, L.A., 1999. Saltatorial leaf-feeding Thysanoptera (Thripidae: Dendrothripinae) from Australia and New Caledonia, with
newly recorded pests of ferns, figs and mulberries. Aust. J.
Entomol. 38, 257273.
Orwig, D.A., Foster, D.R., 1998. Forest response to the introduced
hemlock woolly adelgid in Southern New England. U.S.A. J.
Torrey Bot. Soc. 125, 6073.
Parker, B.L., Skinner, M., Teillon, H.B. (Eds.), 1988. The 1988 thrips
infestation of sugar maple. Proceedings of the regional meeting.
Burlington, Vt., 23 June 1988. Vt. Agric. Exp. Stn. Bull. p. 669.
Parker, B.I., Skinner, M., 1993. Field-evaluation of traps for monitoring emergence of pear thrips (Thysanoptera Thripidae). J.
Econ. Entomol. 86, 4652.

334

S.M. Werner et al. / Forest Ecology and Management 201 (2004) 327334

Parker, G.G., Smith, A.P., Hogan, K.P., 1992. Access to the upper
forest canopy with a large tower crane: sampling the treetops in
three dimensions. BioScience 42, 664671.
Pena, J.E., Duncan, R., 1992. Sampling methods for ProdiplosisLongifila (Diptera Cecidomyiidae) in limes. Environ. Entomol.
21, 9961001.
Raffa, K.F., Hall, D.J., 1989. Thrips calcaratus Uzel (Thysanoptera:
Thripidae), a new pest of basswood trees in the Lake States. Can.
J. For. Res. 18, 16611662.
Raffa, K.F., Hall, D.J., Kearby, W., Katovich, S., 1992. Seasonal life
history of introduced basswood thrips (Thysanoptera: Thripidae)
in Wisconsin, with observations on associated thrips species.
Environ. Entomol. 21, 771779.
Richardson, B.J., Azarbayjani, F.F., Burgin, S., Richardson, S.,
1999. Arboreal arthropod biodiversity in woodlands: effect of
collection procedures and geographic distance on estimates of
diversity found on two species of Melaleuca. Aust. J. Ecol. 24,
544554.
Rieske, L.K., Raffa, K.F., 1996. Bionomics and host range of the
introduced basswood thrips, Thrips calcaratus Uzel, in Wisconsin. Ann. Entomol. Soc. Am. 89, 7580.
SAS Institute, 1999. SAS/StaT Users Guide, Version 8.0. SAS
Institute, Cary, NC.
Schowalter, T.D., 1994. Invertebrate community structure and herbivory in a tropical rainforest canopy in Puerto-Rico following
hurricane Hugo. Biotropica 26, 312319.

Schowalter, T.D., 1995. Canopy arthropod communities in relation

to forest age and alternative harvest practices in western Oregon.
For. Ecol. Manage. 78, 115125.
Shepherd, R.F., Brown, C.E., 1971. Sequential egg band sampling
and probability methods of predicting defoliation by Malacosoma disstria (Lasiocampidae: Lepidoptera). Can. Entomol.
103, 13711379.
Su, J.C., Woods, S.A., 2001. Importance of sampling along a vertical
gradient to compare the insect fauna in managed forests.
Environ. Entomol. 30, 400408.
Stannard, L.J., 1968. The thrips or Thysanoptera of Illinois. Illinois
Nat. History Survey Bull. 29, 215552.
Stork, N.E., 2001. The management implications of canopy
research. Plant Ecol. 153, 313317.
Sweney, W.J., Jones, A.E., 1975. Methods for sampling foliage and
insect populations of the beech forest canopy. N.Z. J. For. Sci. 5,
119122.
USDA FS, 1991. Important Forest Trees of the Eastern United
States. USDA FS FS-466, 111 pp.
Watt, A.D., Stork, N.E., Mcbeath, C., Lawson, G.L., 1997. Impact of
forest management on insect abundance and damage in a lowland tropical forest in Southern Cameroon. J. Appl. Ecol. 34,
985998.
Zandt, H.S., 1994. A comparison of three sampling techniques to
estimate population size of caterpillars in trees. Oecologia 97,
399406.