Agriculture, Ecosystems and Environment 97 (2003) 51–64

Effects of livestock grazing on rangeland grasshopper

(Orthoptera: Acrididae) abundance
Kevin M. O’Neill a,∗ , Bret E. Olson b , Marni G. Rolston a , Roseann Wallander b ,
Deanna P. Larson a,1 , Catherine E. Seibert c
a Department of Entomology, Montana State University, Bozeman, MT 59717, USA
bDepartment of Animal and Range Sciences, Montana State University, Bozeman, MT 59717, USA
c Department of Plant Sciences and Plant Pathology, Montana State University, Bozeman, MT 59717, USA
Received 8 February 2002; received in revised form 14 February 2003; accepted 4 March 2003

Abstract
Livestock may impact habitat quality for grasshoppers by reducing food availability and by altering microclimate and
potential oviposition sites. A 5-year study was conducted to create consistent grazing impacts on replicated plots and measure
their effects on plant cover, microclimate, and grasshopper abundance. Cattle were used to produce two levels of graz-
ing intensity that were compared to ungrazed controls. Differences in plant cover were greatest immediately after grazing
each summer, grasshopper microhabitats tending to be shadier, cooler, less windy, and more humid in the ungrazed plots.
The grasshopper assemblage included five of the worst pest grasshopper species in North America: Ageneotettix deorum,
Aulocara elliotti, Melanoplus sanguinipes, M. packardii, and Camnula pellucida. Most species had greater abundance on
ungrazed pastures, particularly during the 4–6 weeks after grazing each year. However, A. elliotti was often more abun-
dant in heavily grazed areas early in the year when early instars were present and in late summer when adults were pre-
dominant. There was no strong evidence that the effect of grazing on grasshopper abundance increased over the 5-year
study. At this time, all changes in grasshopper numbers cannot be directly attributed to particular habitat characteristics
that changed after grazing, but the results suggest that grazing management could be used to reduce pest grasshopper
densities.
© 2003 Elsevier Science B.V. All rights reserved.
Keywords: Cattle; Grazing management; Grasshopper assemblages; Microclimate; Pest Management; Montana; USA

1. Introduction vertebrate grazers (Hewitt and Onsager, 1983). In

Grasshoppers are major primary consumers of
grassland vegetation and also sever plant materials
that fall to the ground and become unavailable to
∗ Corresponding author. Tel.: +1-406-994-2333;
fax: +1-406-994-6029.
E-mail address: koneill@montana.edu (K.M. O’Neill).
1 Present address: 1205 East 2500 North, North Logan, UT
84341, USA.
competition with domestic livestock, grasshoppers
may consume more foliage than that allocated to
livestock in grazing programs (Onsager, 2000). Con-
versely, foraging by livestock reduces food available
to grasshoppers. Grazing and trampling may also
modify the microclimate, as well as the physical
structure of vegetation and the soil surface, thereby
influencing the quality of the thermal environment and
the availability of oviposition sites. Thus, appropriate
manipulation of the timing and intensity of livestock

0167-8809/03/$ – see front matter © 2003 Elsevier Science B.V. All rights reserved.
doi:10.1016/S0167-8809(03)00136-1
52 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64
grazing could depress grasshopper populations and
reduce the frequency and severity of pest outbreaks
(Curry, 1994; Onsager, 2000).
The effects of grazing on insects have not been em-
phasized in managing western US rangelands (Quigley
et al., 1989; Vavra et al., 1994; Fuhlendorf and Engle,
2001), although studies show that grazing is often fol-
lowed by changes in grasshopper density (Knutsen and
Campbell, 1974; Holmes et al., 1979; Capinera and
Sechrist, 1982; East and Pottinger, 1983; Jepson-Innes
and Bock, 1989; Quinn and Walgenbach, 1990; Miller
and Onsager, 1991; Welch et al., 1991; Fielding and
Brusven, 1993; Onsager, 2000). These studies varied
in methodology, however, some being unreplicated,
some providing no data on individual species or tem-
poral variation in grazing effects. To further investigate
the effect of grazing, a 5-year study was conducted
in southwest Montana, USA, to create two levels of
grazing impacts on replicated plots, measure grasshop-
per abundance multiple times before and after graz-
ing each year, and identify grasshoppers to species. To
gain insight into the mechanisms linking grazing to
grasshopper abundance, the effects of grazing on plant
cover, temperature, wind speed, and humidity were
measured. The objectives were to determine the tim-
ing and magnitude of grasshopper responses to grazing
and variation in species responses. The results indicate
that grazing can cause major changes in grasshopper
microhabitat and that most grasshoppers at the site are
affected negatively by these changes.
2. Materials and methods
The effects of livestock grazing on grasshoppers
was studied 9 km south of Three Forks, Montana,
USA, in a crested wheatgrass (Agropyron cristatum
(L.) Gaertn) pasture that also contained scattered na-
tive grasses and forbs, including needle-and-thread
grass (Stipa comata Trin. and Rupr.), blue gramma
grass (Bouteloua gracilis (H.B.K.) Lag. ex Steud.),
scarlet globemallow (Spharalcea coccinea (Pursh)
Rydb.), and yellow sweetclover (Melilotus officinalis
(L.) Lam.), the latter being a biennial that appeared
in great quantity only in 1993, 1995, and 1997.
Small plots were used so that each would have rel-
atively homogeneous topography and vegetation con-
sisting primarily of A. cristatum rather than Stipa or
Bouteloua. The plots were small enough that changes
in grasshopper abundance would at least partially re-
flect habitat selection by grasshoppers moving into and
out of the plots. In 1993, four 0.37 ha (105 m × 35 m)
blocks were subdivided into three 35 m × 35 m plots
surrounded by temporary electric fencing. Each plot
was assigned to one of three treatments: (1) control—
no grazing from 1993 to 1997, (2) mosaic—grazing
that resulted in a mosaic of grazed and ungrazed plants
(mean ± S.E. = 2.4 ± 0.2 days of grazing per year;
N = 20 = 5 years × 4 plots/year; Table 1), and (3)
uniform—grazing of longer duration that resulted in
uniformly short plants (mean ± S.E. = 4.9 ± 0.3 days;
N = 20; Wilcoxon paired signed-rank test comparing
mosaic to uniform, P < 0.001). The uniform treat-
ments closely mimicked a common grazing system in
North America referred to as “short-duration grazing”
(Holechek et al., 1989). Within a block, the center
plot served as the control and the two others were as-
signed randomly to mosaic and uniform treatments.
Each block was separated from the others by at least
200 m and each plot was assigned to the same grazing
treatment during all 5 years. Cattle were allowed to
graze until the remaining vegetation was judged (visu-
ally) to be at the desired height and patchiness. In June
1993, the electric fences were temporarily opened to
allow access of about 300 cattle in the paddock to uni-
form plots for 7 days and to mosaic plots for 1 day
(with the exception of Block 1 which the cattle did
not enter at first). From 1994 to 1997, grazing was
controlled more precisely by simultaneously enclos-
ing four cattle within each uniform plot for 4–7 days
and each mosaic plot for 2–3 days. Thus, from 1994
to 1997, both treatments were grazed at an intensity
of 10.8 cattle/ha, though for different durations each
year. All grazing occurred during the part of the sea-
son in which the pastures are usually grazed.
Once before and twice after grazing each year
(Table 1), forage structural diversity was assessed non-
destructively with a LiCor LAI-2000 Plant Canopy
Analyzer that measures leaf area index (LAI) of the
vegetation, providing an accurate estimate of standing
plant biomass in crested wheatgrass pastures (Olson
et al., 2000). Within each plot, forage structure was
measured at the four main compass points on each
of three randomly placed 1 m diameter circles; per-
manent markers allowed measurement of LAI in the
same locations each year. The LAI data were analyzed
 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64 53

Table 1
Dates on which measurements and grazing took place during the 5 years of study, in Montana (USA)
Year LAI measurement Grazing

Pre-grazing Post-grazing 1 Post-grazing 2 Date grazing begun Date grazing ended Grazing ended on
on all plots on mosaic plots uniform plots
1993 27 May 3 June 21 July 27 May 28 Maya , 1 Juneb 3 June
1994 7 June 15 June 21 July 7 June 9 June 11 June
1995 13 June 22 June 3 August 15 June 17 June 19 June
1996 2 July 11 July 24 September 5 July 8 July 10 July
1997 17 June 27 June 29 July 20 June 23 June 24 Junec , 25 Juned
a Blocks 2–4.
b Block 1.
c Blocks 1–3.
d Block 4.

with repeated measures, analyses of variance with
grazing treatment and date as main effects (SAS,
1989; von Ende, 1993).
After the 1995 season, plant litter was collected
from the soil surface in five 0.1 m2 (0.36 m diame-
ter) circles in each of the 12 plots; the circles were
at least 5 m from one another in randomly selected
spaces among clumps of crested wheatgrass. Dry mass
of litter was determined after washing the samples
to remove soil and drying them at 38 ◦ C for 7 days.
The five samples within each plot were combined and
treatments were compared using a Kruskal–Wallis test
for overall comparisons and Student–Newman–Keuls
tests for pairwise comparisons.
Microclimate measurements were made under
cloudless conditions on warm days when grasshop-
pers were active. The relationship between grazing
intensity and temperature was examined using Omega
Close Focus Infrared Pyrometers. The person mea-
suring temperature walked 10 parallel, equidistant
transects along the 105 m length of each block, first
traversing a uniform plot, then a control plot, and
finally a mosaic plot; on the next transect, the order
was reversed. The pyrometer was directed downwards
and perpendicular to the soil surface from a height of
60 cm, so that it measured the temperature of a 0.6 cm
diameter spot on the soil surface or a spot up to 1.0 cm
diameter on plant surfaces. On each transect, the py-
rometer made about 30 temperature measurements
producing a single mean temperature that provided
an integrated assessment of the temperatures of vege-
tation and soil surfaces. Data for each block were an-
alyzed separately using Wilcoxon paired signed-rank
tests to compare treatment with control plots; pairs of
data consisted of measurements taken along the same
transect. Measurements were made once before graz-
ing in 1993, 1994, and 1996, twice after grazing in
1993 and 1994, and once after grazing in 1996. Wind
speeds were measured using a Cole-Parmer Tri-Sense
air velocity meter, with an 8 mm diameter sensor. Two
days after grazing in 1993, wind speed was measured
at three heights (0.5, 10, and 50 cm) in 100 randomly
chosen locations within each treatment of Block 3.
Log transformed data were analyzed in a 3 × 3 facto-
rial (grazing treatment × height) analysis of variance.
Relative humidity at the soil surface was measured
using a Cole-Parmer Tri-Sense humidity meter and an
11 mm diameter probe. Humidity measurements were
made after grazing in the control and uniform plots
of Block 3 in 1993 and Blocks 1 and 2 in 1994. Data
were analyzed using Wilcoxon pair signed-rank tests.
Community composition and abundance of
grasshoppers was estimated from 38 cm diameter
sweep net samples, which provide fairly accurate
estimates of the relative abundance of grasshoppers
(Mulkern et al., 1978; Evans et al., 1983; Larson
et al., 1999). Sampling within each plot consisted
of 50 (1993), 100 (1994), or 150 (1995–1997) 180◦
sweeps along an S-shaped transect, parallel to and
10 cm above the ground, and at least 5 m from the
plot boundary to minimize edge effects. Each year,
plots were sampled 1–6 times before grazing and
4–10 times after grazing. For 1993–1995, pre-grazing
counts were analyzed with one-way analyses of vari-
ance. For pre-grazing samples in 1996–1997 and for
post-grazing samples in each of the 5 years, counts
54 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64

were analyzed with repeated measures analyses of

variance with grazing as the between-treatment factor
and date as the within-treatment factor. All counts
were square-root transformed before analysis (Zar,
1996).
After the summers of 1995–1997, densities of
Aulocara elliotti Thomas egg pods were estimated
in control and uniform plots. Each year, twenty
15 cm × 10 cm × 4 cm deep soil samples were col-
lected along transects at least 5 m from plot bound-
aries. At 1 m intervals, soil was excavated from the
nearest patch of bare soil large enough to hold a sam-
pling frame placed against the southeast edge of the
surrounding vegetation, the preferred oviposition site
of this species (Fisher, 1993). After being counted,
pods were placed at a constant 25 ◦ C. Each of the
hundreds of nymphs that emerged was A. elliotti.
Data (square root of egg pod counts) were analyzed
with repeated measures analyses of variance with
grazing as the between-treatment factor and year as
the within-treatment factor.

3. Results

Before grazing in 1993, mean LAI values were sim-

ilar in plots planned for different treatments, but af-
ter grazing LAI diverged widely (Fig. 1). Similarly,
from 1994 to 1997, there were no pre-grazing differ-
ences among treatments. After grazing, control plots
had the highest LAI and uniform treatments the low-
est (except for 1997 when mosaic and uniform treat-
ments did not differ). By late August, LAI values
among the treatments converged, although they re-
mained higher in control than in uniform plots. As the
study progressed, patches of soil among grass clumps
in the control plots accumulated plant litter. By the fall
1995, the amount of plant litter differed among treat-
ments (Kruskal–Wallis test, P = 0.001). Control plots
(mean ±S.E. litter mass = 71.2±5.4 g per 0.5 m2 ) ac-
cumulated more litter than mosaic (44.2 ± 4.8 g) and Fig. 1. Mean LAI values for pre- and post-grazing samples from
uniform plots (29.2 ± 6.9 g) (Student–Newman–Keuls 1993 to 1997 (N = 4). F-values are for treatment×date interactions
tests; P < 0.05 in both comparisons); mosaic and uni- (d.f. = 4, 12), repeated measures analyses of variance (∗ P < 0.01,
∗∗ P < 0.001); means associated with different letter for each date
form treatments did not differ (P > 0.05).
Grazing influenced the microclimate experienced are significantly different (least-square mean tests at α = 0.05
level).
by grasshoppers. Temperatures before grazing in
1993 were similar among treatments. Immediately
after grazing, however, temperatures in control plots
 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64 55

Fig. 2. Differences in mean temperatures between pre- and post-grazing samples (∗ : significantly different at P < 0.05, Wilcoxon paired
signed-rank tests; N = 10).

were lower than in uniform plots in all four blocks
and lower than in the mosaic plots in two of the
blocks (Fig. 2). The differences persisted into late
summer of 1993 and the pre-grazing periods of 1994
(when mosaic temperatures exceeded controls in two
blocks). After grazing in 1994, temperatures were
lowest in controls during June and July. In 1996,
pre-grazing measurements suggested an even stronger
cumulative effect, and post-grazing differences were
again observed. Analysis of wind velocities revealed
a significant treatment × height interaction (F4,891 =
8.82; P < 0.001; N = 900), with the greatest dif-
ferences occurring at 10 cm (Fig. 3). Relative hu-
midity was higher in control than in uniform plots
in all three sets of measurements (Wilcoxon pair
signed-rank tests; 1993, P < 0.01, N = 37; 1994,
P < 0.001, N = 40 (Block 1), P < 0.001, N = 40
(Block 2)).
Pre- and post-grazing sweep samples combined
contained 24 species of grasshoppers, 90% of which
were seven species whose relative proportions var-
ied among years (Fig. 4). The 81,023 grasshoppers
were collected in 54 samples in each of the twelve
1225 m2 plots, so sweep netting removed an average
of 0.10 grasshoppers/m2 /sample; the largest sample
(817 grasshoppers) removed 0.67 grasshoppers/m2 .
Thus, sampling removed a relatively small num-
ber of grasshoppers compared to the standard eco-
nomic threshold for western US rangeland of about
10 grasshoppers/m2 (Skold and Davis, 1996).
Before grazing in 1993, there was no differ-
ence among treatments in the overall abundance of
56 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64

Fig. 3. Mean air velocity at different heights, Block 3 (1993) and

mean relative humidity at the soil surface (N = 100).

grasshoppers (Table 2, Fig. 5). After grazing in 1993,

total abundance continued to be similar among treat-
ments. Pre-grazing samples in 1994 and 1995 also
showed no differences among grazing treatments, but
after grazing, abundance in controls exceeded that in
the other treatments until late July each year; counts
in mosaic plots were intermediate between those in
control and uniform plots. During the pre-grazing Fig. 4. The proportion of each of the seven most abundant species
periods of 1996 and 1997, grasshopper abundance of grasshopper (pre- and post-grazing samples combined); dashed
tended to start out higher in the uniform plots. Af- lines are each species average proportion in combined 5-year
ter grazing, numbers in mosaic and uniform plots counts.
immediately declined, while they remained high in
controls, before declining over several weeks.
During 1993, there was no evidence that Melanoplus was most abundant on control plots and least abun-
sanguinipes responded to grazing (Table 2, Fig. 6). dant on uniform plots in the month after grazing. The
However, there were significant grazing treatment × effect was not strictly related to the stages of M. san-
date interactions in 1994–1997 when M. sanguinipes guinipes present, because the instars most abundant
 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64 57

Table 2
Analyses of variance of pre-grazing (Pre) and post-grazing (Post) grasshopper abundance
Year Period Species Treatment effectsa Treatment × date interactions

F d.f. Probability F d.f. Probability

1993 Pre All species combined 0.35 2, 9 ns – – –

Post 0.45 2, 9 ns 1.33 6, 27 ns
1994 Pre 1.66 2, 9 ns – – –
Post 1.97 2, 9 ns 2.94 16, 72 P < 0.05
1995 Pre 2.56 2, 9 ns – – –
Post 4.59 2, 9 P < 0.05 7.69 18, 81 P < 0.001
1996 Pre 2.10 2, 9 ns 1.82 10, 45 ns
Post 33.41 2, 9 P < 0.001 14.32 16, 72 P < 0.001
1997 Pre 6.19 2, 9 P < 0.05 4.23 6, 27 P < 0.01
Post 0.34 2, 9 ns 7.18 16, 72 P < 0.001
1994 Post A. deorum 0.10 2, 9 ns 1.27 16, 72 ns
1995 Post 0.43 2, 9 ns 2.30 18, 81 P < 0.05
1996 Pre 1.71 2, 9 ns 1.00 10, 45 ns
Post 15.87 2, 9 P < 0.01 8.64 16, 72 P < 0.001
1997 Pre 1.68 2, 9 ns 1.88 6, 27 ns
Post 0.92 2, 9 ns 4.38 16, 72 P < 0.01
1993 Pre A. elliotti 0.26 2, 9 ns – – –
Post 2.33 2, 9 ns 1.58 6, 27 ns
1994 Pre 2.26 2, 9 ns – – –
Post 2.83 2, 9 ns 1.75 16, 72 ns
1995 Pre 9.51 2, 9 P < 0.01 – – –
Post 18.00 2, 9 P < 0.001 3.52 18, 81 P < 0.01
1996 Pre 13.8 2, 9 P < 0.01 7.93 10, 45 P < 0.001
Post 0.05 2, 9 ns 1.48 16, 72 ns
1997 Pre 11.99 2, 9 P < 0.01 2.41 6, 27 ns
Post 2, 90 2, 9 ns 4.26 16, 72 P < 0.01
1995 Post C. pellucida 0.01 2, 9 ns 1.94 18, 81 P < 0.05
1996 Pre 0.43 2, 9 ns 2.16 10, 45 P < 0.05
Post 2.74 2, 9 ns 2.45 16, 72 P < 0.01
1997 Pre 2.00 2, 9 ns 2.05 6, 27 ns
Post 0.49 2, 9 ns 2.49 16, 72 P < 0.01
1994 Post M. infantilis 0.55 2, 9 ns 1.33 16, 72 ns
1995 Post 0.47 2, 9 ns 3.41 18, 81 P < 0.01
1996 Pre 0.08 2, 9 ns 1.91 10, 45 ns
Post 3.45 2, 9 ns 4.65 16, 72 P < 0.001
1997 Pre 0.52 2, 9 ns 1.5 6, 27 ns
Post 0.32 2, 9 ns 4.78 16, 72 P < 0.01
1994 Post M. packardii 0.54 2, 9 ns 1.51 16, 72 0.2
1995 Post 1.54 2, 9 ns 3.3 18, 81 P < 0.05
58 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64

Table 2 (Continued )
Year Period Species Treatment effectsa Treatment × date interactions

F d.f. Probability F d.f. Probability

1996 Pre 2.38 2, 9 ns 1.8 10, 45 ns

Post 12.89 2, 9 P < 0.01 4.82 16, 72 P < 0.01
1997 Pre 5.18 2, 9 P < 0.05 2.74 6, 27 P < 0.05
Post 2.22 2, 9 ns 4.75 16, 72 P < 0.001
1993 Pre M. sanguinipes 0.48 2, 9 ns – – –
Post 0.69 2, 9 ns 0.84 6, 27 ns
1994 Pre 5.64 2, 9 0.03 – – –
Post 6.58 2, 9 0.017 3.43 16, 72 0.016
1995 Pre 0.97 2, 9 ns – – –
Post 35.49 2, 9 P < 0.001 15.83 18, 81 P < 0.001
1996 Pre 22.25 2, 9 P < 0.001 5.86 10, 45 P < 0.001
Post 46.95 2, 9 P < 0.001 13.47 16, 72 P < 0.001
1997 Pre 0.37 2, 9 ns 3.9 6, 27 P < 0.01
Post 10.88 2, 9 P < 0.01 8.92 16, 72 P < 0.001
1994 Post P. nebrascensis 15.65 2, 9 0.0012 2.69 16, 72 P < 0.05
1995 Post 10.49 2, 9 P < 0.01 5.68 18, 81 P < 0.001
1994 Post P. delicatula 1.53 2, 9 ns 1.44 16, 72 ns
1995 Post 2.40 2, 9 ns 1.17 18, 81 ns
a Repeated measures analyses of variance, except for 1993–1995 pre-grazing samples where one-way analysis of variance was used;

ns: not significant at α > 0.05.

in the first sample after grazing varied among years:
1994 (II–III), 1995 (I–II), 1996 (III–V), and 1997
(III). The effect of grazing appeared to carry over
from year to year, as M. sanguinipes was more abun-
dant in control plots before grazing in 1994, 1996,
and 1997. Melanoplus packardii (1995–1997) (Fig. 7)
and Melanoplus infantilis Scudder (1996–1997) re-
sponded to grazing in a manner similar to M. san-
guinipes (Table 2). During the pre-grazing periods
of 1996 and 1997, M. packardii and M. infantilis
abundances in control plots were comparable to or
lower than those in the mosaic and uniform plots.
Immediately after grazing, however, their abundances
stabilized or increased on the control plots, but de-
clined on mosaic and uniform plots. From 1995 to
1997, the response of Ageneotettix deorum (Scudder)
(Fig. 8) and Camnula pellucida Scudder to grazing
was similar to that of Melanoplus (Table 2). In 1994
and 1995 Psoloessa delicatula Scudder exhibited no
difference among treatments, whereas Phoetaliotes
nebrascensis (Thomas) tended to be more abundant
in controls (Table 2).
A. elliotti abundance tended to be higher in uni-
form plots, especially before grazing and in late sum-
mer (Fig. 9). In 1993, A. elliotti was more abundant
on uniform plots in the first samples after grazing. By
late summer 1994 and during the pre-grazing period
of 1995, it was again more abundant on uniform plots.
A similar pattern occurred in late 1995 and, to a lesser
extent, during the pre-grazing period of 1996 (Table 2).
In 1996 and 1997, dual peaks of abundance in uniform
plots occurred when instars I–III (pre-grazing) and
adults (mid- to late-August) were present. Egg pods of
this species were also more numerous on uniform than
control plots. The mean number of egg pods collected
per uniform plot was 22.0 ± 3.6 in 1995, 16.8 ± 2.8 in
1996, and 18.7 ± 5.3 in 1997; means in control plots
were 0 in 1995, 1.0 ± 0.4 in 1996, and 2.3 ± 1.9 in
1997 (repeated-measures analysis of variance; treat-
ment effect: F1,12 = 67.3, P = 0.0002; year effect:
 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64
Fig. 5. Mean number of grasshoppers per sweep sample of all
species combined, 1993–1997 (N = 4); dashed line is the time at
which grazing occurred.
F2,12 = 0.07, P = 0.93; grazing treatment × date in-
teraction: F2,12 = 1.39, P = 0.29).
4. Discussion
Previous studies have found either lower (Capinera
and Sechrist, 1982; Jepson-Innes and Bock, 1989;
Quinn and Walgenbach, 1990; Welch et al., 1991;
59
Fig. 6. Mean number of M. sanguinipes per sweep sample,
1993–1997 (N = 4); dashed line is the time at which grazing
occurred.
Onsager, 2000) or higher (Holmes et al., 1979;
Campbell et al., 1974; Knutsen and Campbell, 1974;
van Wingerden et al., 1991) overall grasshopper
densities following grazing. Within a grasshopper
assemblage, however, species may respond differ-
ently to grazing. Holmes et al. (1979) found that the
abundances of Chortippus longicornis (Latr.) and M.
sanguinipes tended to decrease with grazing intensity,
60 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64
Fig. 7. Mean number of M. packardii per sweep sample, 1995–1997
(N = 4); dashed line is the time at which grazing occurred.
whereas those of C. pellucida, M. dawsoni, and M.
infantilis increased.
In the present study, a reduction in grasshopper
density with increased grazing intensity was observed
for grasshopper assemblage as a whole, for M. san-
guinipes in 1994–1997, and for M. packardii, M. in-
fantilus, A. deorum, and C. pellucida in 1995–1997.
Differences were most pronounced 4–6 weeks follow-
ing grazing each year. The lack of a grazing effect
on grasshopper abundance in 1993 may be partially
due to the substantial regrowth of vegetation that oc-
curred after heavy rains in June and to low sampling
intensity that year. Although P. nebrascensis was more
abundant in control plots after grazing in 1994 and
1995, results for this species are unreliable because its
abundance tends to be overestimated in sweep samples
from ungrazed areas (Larson et al., 1999). The amount
of plant litter accumulating decreased with increasing
grazing intensity, and temperature differences among
treatments carried over from year to year. However,
LAI differences among treatments did not increase af-
ter 1993, and were often similar in the spring each year.
Fig. 8. Mean number of A. deorum per sweep sample, 1995–1997
(N = 4); dashed line is the time at which grazing occurred.
In contrast to the other species, A. elliotti was often
most abundant in the heavily grazed treatments, par-
ticularly in late spring when instars I–III predominated
and in late summer when adults were present. The
greater number of A. elliotti in sweep samples from
uniform plots is consistent with several other observa-
tions. During a behavioral study, it was difficult to find
A. elliotti in control plots, even when it was abundant
on adjacent uniform plots (O’Neill and Rolston, per-
sonal observation). In the late 1980s and early 1990s,
the site was more heavily grazed and A. elliotti was by
far the most common grasshopper, while it remained
relatively rare at an ungrazed crested wheatgrass pas-
ture nearby (O’Neill et al., 1993). Gillespie and Kemp
(1996) found more A. elliotti in grazed pastures than
in adjacent pastures that had been taken out of graz-
ing production. Pfadt (1949) concluded that A. elliotti
was excluded from the dense vegetation of tall grass
prairie.
The grazing treatment × date interactions have sev-
eral possible explanations. Differences in LAI values
 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64
Fig. 9. Mean numbers of A. elliotti per sweep sample, 1993–1997;
N = 4 for each mean; dashed line is the time at which grazing
occurred.
between the grazed and ungrazed plots declined in late
summer, and nitrogen and water content of crested
wheatgrass is often lowest immediately after graz-
ing (Miller and Onsager, 1991). Furthermore, the pe-
riod of greatest vegetational differences among treat-
ments coincided with the presence of nymphs of most
species. During the 4–6 weeks following grazing in
1994–1997, most M. sanguinipes were instars III–V,
but adults predominated from late July to mid-August
when counts in different treatments converged.
61
Individual grasshoppers can move considerable dis-
tances relative to the size the 35 m2 plots in this study.
Joern (1983) found that four species moved an aver-
age of 5–8 m per day. Adult M. sanguinipes moved up
to 90 m per day (Reigert et al., 1954), and grasshopper
nymphs may move tens of meters in less than a week
(Chapman, 1959). As a result, changes in grasshop-
per abundance in the present study probably reflected
not only birth and death rates within plots, but also
migration. A patch with high quality vegetation may
harbor higher grasshopper densities than the surround-
ing habitat (Heidorn and Joern, 1987; Joern, 1996)
which may explain why high densities of sweetclover
in control plots in 1995 were associated with high den-
sities of M. packardii and M. sanguinipes. In 1994,
higher numbers of both species were found in patches
of sweetclover than in adjacent crested wheatgrass at
a nearby site (O’Neill and Larson, unpublished data).
However, green islands of sweetclover existed only
briefly during 3 years, so their existence cannot explain
differences among treatments over the entire study or
even over an entire season. Movement of grasshop-
pers in response to variation in patch quality forms
the basis for using trap strips as a management tool
(Joern, 1996), whereas ungrazed patches of preferred
food plants may exacerbate grasshopper problems on
rangeland planted with legumes (Hewitt et al., 1982),
or where legumes are used as cover crops (Milbraith
et al., 1998).
Ungrazed grassland with dense canopy cover pro-
vides a different microhabitat for grasshoppers than
heavily grazed rangeland (Waterhouse, 1955). Re-
moval of vegetation modifies microclimate by increas-
ing insolation on the soil surface and changing air
flow over the ground (Geiger, 1966; Detling, 1987).
Grasshopper microhabitat on uniform plots tended
to be warmer, windier, drier, and less shaded. Dense
vegetation in ungrazed areas eliminates ground-level
basking sites for some species, whereas heavily grazed
habitats have less shade and fewer elevated perches
for cooling. Microclimate changes may affect hatch-
ing date, early season survival, rates of development,
susceptibility to pathogenic fungi, and availability of
oviposition sites (Parker, 1930; Kemp and Sanchez,
1987; Kemp and Dennis, 1989; Fisher, 1994). Some
species prefer to lay eggs in soil trampled by livestock
(Curry, 1994), whereas A. deorum, C. pellucida, M.
sanguinipes, and M. infantilis oviposit within grass
62 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64
clumps, and A. elliotti oviposits in litter-free patches
of bare soil (Kemp and Sanchez, 1987; Fisher, 1993).
Among the 377 North American grasshopper
species, the six most abundant species in this study
were ranked among the worst pests (Dysart, 1996).
Grasshoppers may compete with livestock, thus hav-
ing a negative economic effect when densities are
above economic thresholds. However, they are also
keystone species in rangeland ecosystems (Quinn
et al., 1993), as well as being food for predators
(Belovsky and Slade, 1993; O’Neill, 1995) and nu-
trient recyclers (O’Neill, 1994; O’Neill et al., 1997;
Woods et al., 1997). Because sweep net data cannot
be reliably converted to densities, changes in abun-
dance after grazing cannot be related to an economic
threshold. Thus, although it is possible that grazing
reduced grasshopper density below the economic
threshold, it is also possible that it reduced it to lev-
els detrimental to long-term rangeland productivity if
densities remained low.
The present results and those of previous studies
(Curry, 1994; Onsager, 2000) suggest that specific gra-
zing management systems have the potential to re-
duce grasshopper populations. However, each local
grasshopper assemblage may respond quite differently
to grazing. For example, C. pellucida and M. infan-
tilis may respond with increased (Holmes et al., 1979)
or decreased densities (present study). Similarly, short
duration grazing could decrease grasshopper density
in assemblages dominated by M. sanguinipes and in-
crease its densities where A. elliotti predominates. Fu-
ture studies should also assess whether the use of graz-
ing management to impact grasshoppers is compati-
ble with other aims of grazing management (East and
Pottinger, 1983; Quigley et al., 1989; Onsager, 1996).
Acknowledgements
We would like to thank John Beaver, Peter Jensen,
Perry Hofferber, Ruth O’Neill, and Jonathan Turnbow
for assistance with field work and sample processing.
Sue Blodgett, William Kemp, Ted Evans, and sev-
eral anonymous reviewers made insightful comments
on the manuscript. This work was supported by the
Montana Agricultural Experiment Station and by two
grants from the USDA-CSREES Rangeland Research
Grants Program.
References
Belovsky, G.E., Slade, J.B., 1993. The role of vertebrate and
invertebrate predators in a grasshopper community. Oikos 68,
193–201.
Campbell, J.B., Arnett, W.H., Lambley, J.D., Jantz, O.K., Knutson,
H., 1974. Grasshoppers (Acrididae) of the Flints Hills native
tallgrass prairie in Kansas. Kansas Agricultural Experimental
Station Research Paper 19.
Capinera, J.L., Sechrist, T.S., 1982. Grasshopper (Acrididae)-host
plant associations: response of grasshopper populations to cattle
grazing intensity. Can. Entomol. 114, 1055–1062.
Chapman, R.F., 1959. Observations on the flight activity of the
red locust, Nomadacris septemfasciata (Serville). Behaviour 14,
300–344.
Curry, J.P., 1994. Grassland Invertebrates: Ecology, Influence on
Soil Fertility, and Effects on Plant Growth. Chapman & Hall,
New York, 437 pp.
Detling, J.K., 1987. Grass response to herbivory. In: Capinera, J.L.
(Ed.), Integrated Pest Management on Rangeland. Westview
Press, Boulder, CO, pp. 56–68.
Dysart, R.J., 1996. Relative importance of rangeland grasshoppers
in western North America: a numerical ranking from the
literature. In: Cunningham, G.L., Sampson, M.W. (Eds.),
Grasshopper Integrated Pest Management User Handbook.
United States Department of Agriculture, Animal and
Plant Health Inspection Service Technical Bulletin 1809,
pp. VI.6-1–VI.6-20.
East, R., Pottinger, R.P., 1983. Use of grazing animals to control
insect pests of pasture. N. Z. Entomol. 7, 352–359.
Evans, E.W., Rogers, R.A., Opferman, D.J., 1983. Sampling
grasshoppers (Orthoptera: Acrididae) on burned and unburned
tallgrass prairie: night trapping vs. sweeping. Environ. Entomol.
12, 1449–1454.
Fielding, D.J., Brusven, M.A., 1993. Grasshopper (Orthoptera:
Acrididae) community composition and ecological disturbance
on southern Idaho rangeland. Environ. Entomol. 22, 71–
81.
Fisher, J.R., 1993. Location of egg pods of Aulocara elliotti
(Orthoptera: Acrididae) in a field of crested wheatgrass in
Montana. J. Kans. Entomol. Soc. 65, 416–420.
Fisher, J.R., 1994. Temperature effect on postdiapause development
and survival of embryos of three species of Melanoplus
(Orthoptera: Acrididae). Ann. Entomol. Soc. Am. 87, 604–608.
Fuhlendorf, S.D., Engle, D.M., 2001. Restoring heterogeneity
on rangelands: ecosystem management based on evolutionary
grazing patterns. Bioscience 51, 625–632.
Geiger, R., 1966. The Climate Near the Ground. Harvard University
Press, Cambridge, MA, 611 pp.
Gillespie, R.L., Kemp, W.P., 1996. Habitat associations of
grasshopper species (Orthoptera: Acrididae) in winter wheat
(Triticum aestivum L.) and adjacent rangeland. J. Kans.
Entomol. Soc. 68, 415–424.
Heidorn, T.J., Joern, A., 1987. Feeding responses and spatial
distribution of grasshoppers (Acrididae) in response to nitrogen
fertilization of Calamovilfa longifolia. Funct. Ecol. 1, 369–
375.
 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64
Hewitt, G.B., Onsager, J.A., 1983. Control of grasshoppers on
rangeland in the United States: a perspective. J. Range Manage.
36, 202–207.
Hewitt, G.B., Wilton, A.C., Lorenz, R.J., 1982. The suitability of
legumes for rangeland interseeding and as grasshopper food. J.
Range Manage. 35, 653–656.
Holechek, J.L., Pieper, R.D., Herbel, C.H., 1989. Range
Management Principles and Practices. Prentice-Hall,
Englewood Cliffs, NJ, 501 pp.
Holmes, N.D., Smith, D.S., Johnston, A., 1979. Effect of grazing
by cattle on the abundance of grasshoppers on fescue grassland.
J. Range Manage. 32, 310–311.
Jepson-Innes, K., Bock, C.E., 1989. Response of grasshoppers
(Orthoptera: Acrididae) to livestock grazing in southeastern
Arizona: differences between seasons and subfamilies.
Oecologia 78, 430–431.
Joern, A., 1983. Small-scale displacements of grasshoppers
(Orthoptera: Acrididae) within arid grasslands. J. Kans.
Entomol. Soc. 56, 131–137.
Joern, A., 1996. Host plant quality and grasshopper populations.
In: Cunningham, G.L., Sampson, M.W. (Eds.), Grasshopper
Integrated Pest Management User Handbook. United States
Department of Agriculture, Animal and Plant Health Inspection
Service Technical Bulletin 1809, pp. IV.4-1–IV.4-6.
Kemp, W.P., Dennis, B., 1989. Development of two rangeland
grasshoppers at constant temperatures: development thresholds
revisited. Can. Entomol. 121, 363–371.
Kemp, W.P., Sanchez, N.E., 1987. Differences in post-diapause
thermal requirements for eggs of two rangeland grasshoppers.
Can. Entomol. 119, 651–653.
Knutsen, H., Campbell, J.B., Relationships of grasshoppers
(Acrididae) to burning, grazing, and range sites of native
tallgrass prairie in Kansas. In: Proceedings of the Annual
Tall Timbers Conference on Ecology, Animal Control Habitat
Management, vol. 6, pp. 107–120.
Larson, D.P., O’Neill, K.M., Kemp, W.P., 1999. Evaluation of
the accuracy of sweep sampling in determining grasshopper
(Orthoptera: Acrididae) community composition. J. Agric.
Urban Entomol. 16, 207–214.
Milbraith, L.R., Weiss, M.J., Anderson, P.L., Dipirro, M., 1998.
Suitability of legume cover crops for grasshopper (Orthoptera:
Acrididae) development and reproduction. J. Econ. Entomol.
91, 1186–1195.
Miller, R.H., Onsager, J.A., 1991. Grasshopper (Orthoptera:
Acrididae) and plant relationships under different grazing
intensities. Environ. Entomol. 20, 807–814.
Mulkern, G.B., Records, J.C., Carlson, R.B., 1978. Attractants and
phagostimulants used for control and estimation of grasshopper
populations. Ent. Exp. Appl. 24, 350–361.
Olson, B.E., Wallander, R.T., Beaver, J.M., 2000. Comparing
nondestructive measures of forage structure and phytomass.
Can. J. Plant Sci. 80, 565–573.
O’Neill, K.M., 1994. Livestock dung as a food resource and
thermal refuge for rangeland grasshoppers. Pan-Pac. Entomol.
70, 222–229.
O’Neill, K.M., 1995. Digger wasps (Hymenoptera: Sphecidae) and
robber flies (Diptera: Asilidae) as predators of grasshoppers
63
(Orthoptera: Acrididae) on Montana rangeland. Pan-Pac.
Entomol. 71, 248–250.
O’Neill, K.M., Woods, S.A., Streett, D., O’Neill, R.P., 1993.
Aggressive interactions and feeding success in scavenging
grasshoppers (Orthoptera: Acrididae). Environ. Entomol. 22,
751–758.
O’Neill, K.M., Woods, S.A., Streett, D., 1997. Grasshopper
(Orthoptera: Acrididae) foraging on grasshopper feces:
observational and rubidium-labeling studies. Environ. Entomol.
26, 1224–1231.
Onsager, J.A., 1996. The importance of grazing strategies to
grasshopper management: an introduction. In: Cunningham,
G.L., Sampson, M.W. (Eds.), Grasshopper Integrated Pest
Management User Handbook. United States Department of
Agriculture, Animal and Plant Health Inspection Service
Technical Bulletin 1809, pp. V.1-i–V.
Onsager, J.A., 2000. Suppression of grasshoppers in the Great
Plains through grazing management. J. Range Manage. 53,
592–602.
Parker, J.R., 1930. Some effects of temperature and moisture
upon Melanoplus mexicanus mexicanus Saussure and Camnula
pellucida Scudder (Orthoptera), Mont. Agric. Exp. Sta. Bull.
223.
Pfadt, R.E., 1949. Food plants, distribution and abundance of
the big-headed grasshopper, Aulocara elliotti (Thos.). J. Kans.
Entomol. Soc. 22, 69–74.
Quigley, T.M., Dillard, D.S., Reese, J.B., Free, J.C., Henke,
G., Wasser A.S., Feakes, N., 1989. New criteria for
measuring range management activities. USDA For. Serv. GTR-
248.
Quinn, M.A., Walgenbach, D.D., 1990. Influence of grazing history
on the community structure of grasshoppers of a mixed-grass
prairie. Environ. Entomol. 19, 1756–1766.
Quinn, M.A., Johnson, P.S., Butterflied, C.H., Walgenbach, D.D.,
1993. Effect of grasshopper (Orthoptera: Acrididae) density
and plant composition on growth and destruction of grasses.
Environ. Entomol. 22, 993–1002.
Reigert, P.W., Fuller, R.A., Putnam, L.G., 1954. Studies
on dispersal of grasshoppers (Acrididae) tagged with
phosphorus-32. Can. Entomol. 86, 223–232.
SAS, 1989. SAS/STAT User’s Guide, version 6, vol. 2, 4th ed.
SAS Institute Inc., Cary, NC, 1028 pp.
Skold, M.D., Davis, R.M., 1996. Applying economics to
grasshopper management. In: Cunningham, G.L., Sampson,
M.W. (Eds.), Grasshopper Integrated Pest Management User
Handbook. United States Department of Agriculture, Animal
and Plant Health Inspection Service Technical Bulletin 1809,
pp. VI.3-1–VI.3-4.
van Wingerden, W.K.R.E., Musters, J.C.M., Kleukers, R.M.J.C.,
Bongers, W., van Biezen, J.B., 1991. The influence of
cattle grazing intensity on grasshopper abundance (Orthoptera:
Acrididae). Proc. Exp. Appl. Entomol. 2, 28–34.
Vavra, M., Laycock, W.A., Pieper, R.D., 1994. Ecological
Implications of Livestock Herbivory in the West. Society for
Range Management, Denver.
von Ende, C.N., 1993. Repeated measures analysis: growth and
other time-dependent measures. In: Scheiner, S.M., Gurevitch,
64 K.M. O’Neill et al. / Agriculture, Ecosystems and Environment 97 (2003) 51–64

J. (Eds.), Design and Analysis of Ecological Experiments.
Chapman & Hall, New York, pp. 113–137.
Waterhouse, F.L., 1955. Microclimatological profiles in grass cover
in relation to biological problems. Quart. J. Roy. Meteorol. Soc.
81, 63–71.
Welch, J.L., Redak, R., Kondratieff, B.C., 1991. Effect of cattle
grazing on the density and species of grasshoppers (Orthoptera:
Acrididae) of the Central Great Plains experimental range,
Colorado: a reassessment after two decades. J. Kans. Entomol.
Soc. 64, 337–343.
Woods, S.A., O’Neill, K.M., Streett, D.A., 1997. Scavenging
behavior of rangeland grasshoppers (Orthoptera: Acrididae):
rubidium-marker studies. Environ. Entomol. 26, 789–
796.
Zar, J.H., 1996. Biostatistical Analysis, 3rd ed. Prentice-Hall,
Upper Saddle River, NJ, 718 pp.