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Great Plains Research: A Journal of Natural and
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10-1-2009
THE GSSHOPPERS ARPHIA
XANTHOPTE AND DICHROMORPHA
VIRIDIS PREFER INTRODUCED SMOOTH
BROME OVER OTHER GSSES
Sean D. Whipple
University of Nebraska at Kearney
Mathew L. Brust
University of Nebraska-Lincoln
Wya Hoback
University of Nebraska at Kearney, hobackww@unk.edu
Kerri M. Farnsworth-Hoback
University of Nebraska at Kearney
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Whipple, Sean D.; Brust, Mathew L.; Hoback, Wya; and Farnsworth-Hoback, Kerri M., "THE GSSHOPPERS ARPHIA
XANTHOPTE AND DICHROMORPHA VIRIDIS PREFER INTRODUCED SMOOTH BROME OVER OTHER GSSES"
(2009). Great Plains Research: A Journal of Natural and Social Sciences. Paper 1067.
hp://digitalcommons.unl.edu/greatplainsresearch/1067
Manuscript received for review, March 2009; accepted for publication,
June 2009.
THE GRASSHOPPERS ARPHIA XANTHOPTERA AND
DICHROMORPHA VIRIDIS PREFER INTRODUCED SMOOTH
BROME OVER OTHER GRASSES
Sean D. Whipple
Department of Biolog y
University of Nebraska at Kearney
905 West 25th Street
Kearney, NE 68849
Mathew L. Brust1
Department of Entomology
University of Nebraska–Lincoln
202 Plant Industry Building
Lincoln, NE 68583
W. Wyatt Hoback and Kerri M. Farnsworth-Hoback
Department of Biolog y
University of Nebraska at Kearney
905 West 25th Street
Kearney, NE 68849
hobackww@unk.edu
Key Wo rd s: feeding preference, grasshopper, non-native grass, smooth brome, tallgrass prairie
AB STR ACT—A study of feeding preference was conducted on two tallgrass prairie grasshopper species, the
autumn yellow-winged grasshopper Arphia xanthoptera (Burmeister) and the short-winged green grasshopper
Dichromorpha viridis (Scudder), to determine if they would feed upon introduced grass species. Both grasshop-
pers were offered two non-native cool-season grasses, smooth brome (Bromus inermis Leyss) and Kentucky
bluegrass (Poa pratensis L.), and two native warm-season grasses, big bluestem (Adropogon gerardii Vitman)
and sideoats grama (Bouteloua curtipendula Michx.). Live biomass of the plants was weighed before and after
feeding to quantify the amount of each plant species consumed by the grasshoppers. Statistical analysis showed
that D. viridis strongly preferred smooth brome (P ≤ 0.05) over other species offered. A. xanthoptera also con-
sumed more smooth brome than the other grass species offered. These results suggest that both grasshopper
species accept non-native grasses and perhaps prefer them to tallgrass prairie species. Because the tallgrass
prairie ecosystem of the Great Plains has been dramatically impacted by human activity, documentation of the
response of native insects to incursion by exotic plants is important to preservation efforts. Moreover, if grass-
hoppers feed on invasive sod-forming species such as smooth brome and Kentucky bluegrass, they may become
an important ally in maintaining native plant diversity in remnant grassland ecosystems.
INTRODUCTION
The tallgrass prairie is a highly altered ecosystem of
which less than 1% of the original prairie remains (Cully
et al. 2003). Among impacts to this once vast ecosystem,
human alterations include changes in land use, grazing
regimes, re regimes, and plant community composition
(Samson and Knopf 1994). The intentional and accidental
introductions of non-native species have further altered the
ecosystem and increased pressures on the remaining prai-
rie species. Among invertebrates shown to be impacted by
changes to the tallgrass prairie ecosystem, much focus has
been placed on butteries, true bugs, and ground beetles
(e.g., Arenz 1995; Swengel and Swengel, 1998). Less atten-
tion has focused on grasshoppers (Orthoptera: Acrididae)
1Current address: Department of Biology, Chadron State Col-
lege, 1000 Main Street, Chadron, NE 69337
179
Great Plains Research 19 (Fall 2009):179-86
© 2009 Copyright by the Center for Great Plains Studies, University of Nebraska–Lincoln
Great Plains Research Vol. 19 No. 2, 2009180
© 2009 Center for Great Plain s Studies , University of Nebraska–L incoln
because most species are generalist herbivores and thus (it
is assumed), less likely to be impacted by changes to the
plant community.
There are over 400 species of short-horned grasshop-
pers in the United States (Lockwood 2001), of which
more than 100 occur in Nebraska (Brust et al. 2008).
Short-horned grasshoppers (family Acrididae) are im-
portant because they are litter producers, valuable food
for birds and other wildlife, and some species act as weed
control agents on plants that are not palatable to live-
stock (Parker 1984). Some grasshoppers are also of sig-
nicant economic importance as serious rangeland pests
(Mulkern et al. 1969). Grasshopper species composition
is largely determined by food plants and microhabitat
characteristics (Joern and Lawlor 1981), and species may
have narrow requirements for survival (Haarstad 1990;
Ballard and Greenlee 1996; Reed 1996).
Most grasshoppers feed on a range of plants (Joern
1983) and as such are considered generalists with respect
to diet breadth. However, grasshoppers are not indifferent
feeders and often select plants with specic characteris-
tics to gain required nutrients such as nitrogen (Behmer
and Joern 1993). However, plants also contain physical
and chemical defenses against herbivores. Specialist her-
bivores have evolved the ability to tolerate or overcome
these defenses, which are effective in deterring general-
ists (species that consume a variety of food plants). Recent
studies have shown that generalist grasshoppers respond
to changes in plant species composition at the community
level (Stoner and Joern 2004), and grasshopper herbivory
may play a role in shaping the relative abundance of na-
tive and non-native plant species in invaded communities
(Branson and Sword 2008). However, the response of na-
tive generalist insect herbivores such as grasshoppers to
non-native plants has not often been quantied.
Smooth brome (Bromus inermis L.) is one of many
non-native species that h ave become abunda nt in the Gre at
Plains (Whitmore 2000) and remaining tallgrass prairie
(Willson and Stubbendieck 1996). Kentucky bluegrass
has been repeatedly introduced since the early 1800s and
has become naturalized throughout much of the United
States; it is listed as an invasive weed in the Great Plains
states (Wenneberg 2004). Like other non-native grass
species that occur in tallgrass prairies, smooth brome and
Kentucky bluegrass are cool-season (C3) plants (Cully et
al. 2003). Their physiology and phenology is much differ-
ent from the native prairie ora, which is dominated by
warm-season (C4) species. Smooth brome and Kentucky
bluegrass grow actively during the fall and early spring,
owering in late spring or early summer; by midsummer,
seed is mature (Howard 1996; Uchytil 1993). Kentucky
bluegrass becomes nearly dormant during the midsum-
mer while smooth brome will continue growing as long
as moisture is available (Howard 1996; Uchytil 1993).
Big bluestem does not begin actively growing in spring
until several weeks after the cool-season grasses have
begun green-up, owering between July and September
in Nebraska (Uchytil 1988). The timing of active growth
and owering of sideoats grama is very similar to that of
big bluestem (Wasser 1982).
The incursion of smooth brome and other non-native
cool-season plants in Nebraska is likely to have affected
food availability for many tallgrass prairie herbivore
species (Ogle et al. 2003). Porter and Redak (1997)
demonstrated that Melanoplus sanguinipes Fabricius
preferred native grasses over introduced grasses, al-
though forbs comprised the largest proportion of their
diet. Other prairie grasshopper species, including the
autumn yellow-winged grasshopper Arphia xanthoptera
(Burmeister) and the short-winged green grasshopper
Dichromorpha viridis (Scudder), are also likely to have
been impacted.
These two grasshopper species were chosen for study
because little is known of their feeding preferences.
Although D. viridis and A. xanthoptera occur through-
out most of the eastern United States, both species are
thought to be tallgrass prairie specialists (Bragg 1939;
Wilbur and Fritz 1940; Reed 1996). We hypothesized that
these species would prefer native warm-season grasses
over non-native cool-season grasses. However, we found
both grasshopper species to select non-native grasses in
laboratory tests. Our results provide insight into response
of native insect herbivores to non-native species and to the
potential herbivore responses to non-native grass species
on a dwindling tallgrass prairie ecosystem.
MATERIALS AND METHODS
Our metho ds were similar to those of Gang were (1961),
who observed grasshopper feeding damage on a range of
test plants. However, we also collected quantitative data
by weighing plant biomass before and after feeding tri-
als. Adults of the green grasshopper and A. xanthoptera
were collected from unmowed grassy areas using sweep
nets at Lincoln Wilderness Park, Lancaster County, NE
(4513667 N, 692758 E). Specimens were collected in late
August of 2006 and 2007. After collection, grasshoppers
were transported to the University of Nebraska at Kear-
ney where each species was stored separately in plastic
containers with plants from the collection site. Specimens
The Grasshoppers Arphia xanthoptera and Dichromorpha viridis • Sean D. Whipple et al. 181
© 2009 Center for Great Plain s Studies , University of Ne braska–L incoln
were stored at room temperature (approximately 25°C)
for two days prior to the feeding trials.
The day before each feeding trial was conducted,
samples of four plant species were collected at Cottonmill
Park and Recreation Area on the Oldfather Prairie Re-
serve, 2.4 km west of Kearney, NE. The four plant species
chosen were smooth brome (Bromus inermis Leyss.), big
bluestem (Adropogon gerardii Vitman), sideoats grama
(Bouteloua curtipendula Michx.), and Kentucky blue-
grass (Poa pratensis L.). Specimens of the four plant spe-
cies were selected based on healthy appearance (green,
unwilted). Samples were collected from multiple plants (5
to 10 depending on species) by snipping the base of plants
using garden shears and wrapping vegetation in a moist
paper towel. After collection, the plants were placed in
water overnight to prevent dehydration.
Feeding studies consisted of placing a single grass-
hopper specimen into a container with each of the four
plant species. Prior to feeding studies, sections approxi-
mately 25 cm long from the tip of the stem were prepared.
In 2006 each plant specimen was examined for damage,
weighed to the nearest hundredth of a gram, and placed
into a small cylindrical tube. The sections of grasses
weighed approximately 0.8 g for sideoats grama, 1.3 g
for the big bluestem and Kentucky bluegrass, and 3.0 g
for smooth brome. The tubes were lled with a water-
saturated cotton ball to keep the plants from drying out
during the feeding trial. A steel ball was also placed in
the bottom of the tubes to prevent them from falling over
from the weight of the plant or the grasshoppers feeding
on them. Ten replicates were created for each grasshopper
species, for a total of 20 containers with four plant spe-
cies per container. The containers were then placed into
a growth chamber at 25°C on a 12:12 light/dark cycle.
The containers were left in the growth chamber for three
days to allow for feeding, and then removed to weigh the
plants. Each plant was examined for damage to leaves
from feeding. All remaining material was weighed to
the nearest hundredth of a gram. Any clippings that had
fallen to the bottom of the container were identied by
texture and appearance and were weighed and included
in the totals of mass remaining after feeding.
The experiment was repeated in 2007 with 15 rep-
licates per grasshopper species. As a control for mass
change associated with water uptake, six containers were
prepared in the same manner as the experimental groups,
except that no grasshopper was put in with the plants.
Plants were weighed before placing them in the growth
chamber and after three days. Gains in mass were inter-
preted as water uptake by the plant.
Feeding preference data were analyzed following the
methods Rodrigues et al. (2008). Once data were col-
lected, a nonparametric Kruskal-Wallis one-way analysis
of variance was performed to analyze feeding on the four
plant species by each grasshopper species and by each
year. If signicant differences were detected among the
groups, Dunn’s method of pairwise comparisons was
used to separate the means.
RESULTS
The control showed that no plants suffered mass loss
after three days in the growth chamber. Mean water up-
take for smooth brome was 0.075 g. Sideoats grama and
Kentucky bluegrass also gained mass from water uptake
(0.068 g and 0.093 g, respectively). Big bluestem had a
slightly higher mean level of water uptake at 0.151 g. Total
mass gain was between 3% and 16% across plant species.
These values were not signicantly different across the
four plant species (P = 0.469).
In 2006 the 10 A. xanthoptera consumed a total of
1.18 g of smooth brome compared to 0.10 g of sideoats
grama, 0.22 g of big bluestem, and 0.13 g of Kentucky
bluegrass (Fig. 1). Feeding was evident on all the samples
of smooth brome, while in many cases the other three spe-
cies of plant were not fed upon. Grasshoppers ate signi-
cantly more smooth brome (P ≤ 0.05) than sideoats grama
or Kentucky bluegrass. Although grasshoppers ate more
brome grass than big bluestem (Fig. 1), the difference
was not signicant (P = 0.076). Individual grasshoppers
ate variable amounts of grasses with one A. xanthoptera
consuming 0.54 g of smooth brome.
In 2007 the 15 A. xanthoptera together consumed a to-
tal of 7.53 g, with smooth brome comprising 64% (4.82 g)
of the total. Kentucky bluegrass, the other non-native
grass, accounted for 2.31 g of the total, while sideoats
grama and big bluestem accounted for 0.21 and 0.20 g,
respectively (Fig. 1). Mean consumption of smooth brome
was 0.321 g per grasshopper with two of the grasshop-
pers tested exhibiting no feeding on smooth brome. A.
xanthoptera showed much less feeding on the other plant
species, with a mean consumption of 0.014 g of sideoats
grama, 0.013 g of big bluestem, and 0.157 g of Kentucky
bluegrass. In 2007 grasshoppers consumed signicantly
more smooth brome (P ≤ 0.001) than other plant species,
with the exception of Kentucky bluegrass (P = 0.280.05)
(Fig. 1), although there was variation among individuals.
D. viridis strongly preferred smooth brome in both years.
In 2006 individuals consumed a total of 2.38 g of smooth
brome, compared to 0.37 g of sideoats grama, 0.30 g of big
Great Plains Research Vol. 19 No. 2, 2009182
© 2009 Center for Great Plain s Studies , University of Nebraska–L incoln
bluestem, and 0.13 g of Kentucky bluegrass, with feeding
evident on all samples of smooth brome ( Fig. 2). In all cases,
D. viridis were feeding on brome when the container was
opened to weigh the plants at the end of the trial. Although
there was more evidence of feeding on the other three plant
species than was observed for A. xanthoptera, consump-
tion of smooth brome was signicantly higher than that of
the three other plant species (P ≤ 0.001; Fig. 2).
In 2007 D. viridis ate a total of 5.89 g of all plants,
with smooth brome representing 83% (4.88 grams) of the
total plant mass consumed, signicantly more (P < 0.001)
than the other tested species (Fig. 2). Kentucky bluegrass
consumption was second highest with a total of 0.63 g.
Sideoats grama and big bluestem consumption totaled
0.22 g and 0.17 g, respectively. Smooth brome was the
only plant species to show evidence of feeding by all
individual grasshoppers.
The number of plant species consumed by each in-
dividual grasshopper was also compared (Table 1). Both
species of grasshopper fed on two plant species in most
of the trials. A. xanthoptera consumed one, two, or three
plants, but never consumed all four. D. viridis sampled all
plants in 28% of trials.
DISCUSSION
Our experiments revealed surprising results: Two
grasshopper species hypothesized to be tallgrass prairie
specialists fed preferentially on non-native smooth brome
and Kentucky bluegrass over grasses readily available
TABLE 1
PERCENTAGE OF TOTAL NUMBER OF GRASSHOPPERS
OF EACH SPECIES (N = 25) FEEDING ON ONE OR MORE
OF THE OFFERED PLANT SPECIES
Number of plant species
fed upon
Grasshopper species 1234
Arphia xanthoptera 20% 44% 36% 0%
Dichromorpha viridis 20% 36% 16% 28%
Figure 1. The mean amount (±1 standard error) of each plant species consumed by the autumn yellow-winged grasshopper,
Arphia xanthoptera, in laboratory feeding trials in 2006 (N = 10) and 2007 (N = 15). Brome = smooth brome (Bromis inermis)
and Bluegrass = Kentucky bluegrass (Poa pratensis) are non-native cool-season species); Sideoats = sideoats grama (Bouteloua
curtipendula) and Big Blue = big bluestem (Adropogon gerardii) are native warm -season grasses.
The Grasshoppers Arphia xanthoptera and Dichromorpha viridis • Sean D. Whipple et al. 183
© 2009 Center for Great Plain s Studies , University of Ne braska–L incoln
in native tallgrass prairie. Both A. xanthoptera and D.
viridis consumed more smooth brome than other grass
species offered. This observation can either be attributed
to the grasshoppers encountering a relatively novel food
or having shifted their diet choices because of the abun-
dance of smooth brome within their community. In our
experiments, we tested non-native cool-season grasses
versus warm-season grasses commonly found in native
tallgrass prairies. Because these grasses are very different
phylo genetically and physiologically, the latter explana-
tion seems most likely. For A. xanthoptera, analysis of the
2006 trials showed a statistical difference between smooth
brome and all other plants tested, except the native big
bluestem. In 2007 smooth brome consumption was sig-
nic a n tly hig h e r than all plants except Kent u ck y blu eg r a ss.
While we cannot know whether A. xanthoptera historically
fed upon native cool-season grasses, our results suggest
that cool-season non-native grasses are acceptable and
even preferred food plants during the late summer.
D. viridis are adults in summer and fall, and eggs hatch
in the spring (Otte 1981). A. xanthoptera adults are present
July through November (Otte 1984). Time of hatching and
maturation may be related to preference for cool-season
versus warm-season grasses. If smooth brome is readily
available and contains the necessary nutrients for survival
and reproduction, it is not surprising that it is utilized
(Ueckert et al. 1972). However, because of defense com-
pounds, it is rare for even generalist herbivores to eat non-
native species, especially when native species are present
(Hierro and Callaway 2003; Zhang and Jiang 2006). Hinks
and Olfert (1999) previously showed nymphal Melanoplus
sanguinipes to survive on smooth brome. However, Olfert
et al. (1994) found that of the plants tested, smooth brome
was the most detrimental and resulted in slower develop-
ment and a lower mean dry weight.
The difference in results may also be a result of dif-
ferent plant nutrient levels, as Joern and Behmer (1997)
demonstrated for Ageneotettix deorum. It is difcult to
predict whether the feeding preferences of grasshoppers
tested in this study would be different if tested earlier in
the season or when they are in the nymphal stage. Future
studies should examine feeding preferences of more
Figure 2. The mean amount (±1 standard error) of each plant species consumed by short-winged green grasshopper, Dichromorpha
viridis, in laboratory feeding trials in 2006 (N = 10) and 2007 (N = 15). Brome = smooth brome (Bromis inermis) and Bluegrass =
Kentucky bluegrass (Poa pratensis) are non-native cool-season species); Sideoats = sideoats grama (Bouteloua curtipendula) and
Big Blue = big bluestem (Adropogon gerardii) are native warm-season grasses.
Great Plains Research Vol. 19 No. 2, 2009184
© 2009 Center for Great Plain s Studies , University of Nebraska–L incoln
grasshopper species with more plant choices to separate
phenology, plant physiology, and nutrient condition.
Various authors have debated whether generalist in-
sect herbivores prefer cool-season or warm-season plant
species (e.g., Boutton et al. 1978; Pinder and Kroh 1987).
Heidorn and Joern (1984) found that Ageneotettix deorum
(Scudder), preferred cool-season grasses over warm-sea-
son grasses. They found no correlation between preference
and leaf water content, crude protein content, or tough-
ness. They concluded that differences in leaf anatomy
between cool-season and warm-season grasses resulted
in the observed preference. Our results lend only partial
support to this assertion. While the food plant chosen by
both grasshopper species was a cool-season grass, the two
cool-season species were not always clearly preferred over
the warm-season grasses offered. For some individuals, the
amount of Kentucky bluegrass consumed was lower than
the amount eaten of either species of warm-season grass.
Although we tested different species of grasses and grass-
hoppers, Heidorn and Joern’s conclusion that leaf anatomy
is a strong in uence on grasshopper feeding preference for
cool-season grasses appears to be invalid.
Behmer and Joern (1993) demonstrated that grasshop-
pers feed based on a need for a limiting nutrient. During
development, grasshoppers are often limited by nitrogen,
which they use for molting, growth, and reproduction
(Joern and Behmer 1997). Because these grasshoppers
are adults late in the season, plants in our study were
collected in late August. The warm-season grasses had
already owered and Kentucky bluegrass may have been
dormant. It is possible that smooth brome was preferred
because it was the only actively growing food plant. How-
ever, the relationship between food preference and leaf
water content has been studied by others, with conict-
ing results. Some studies resulted in little or no correla-
tion between leaf water content and feeding preference
(Gangwere 1961; Heidorn and Joern 1984). Lewis (1984)
showed that Melanoplus differentialis Thomas prefers
to feed on the wilted vegetation of the wild sunower,
Helianthus annuus L., rather than undamaged tissue. If
grasshoppers prefer water-stressed plants, then we would
expect that the plants that took up the most water in our
study to also be those that were preferred. Because mean
water uptake was not signicantly different among the
plants we tested, we cannot resolve this question.
It is difcult to draw conclusions about feeding choices
from the percentage of individual grasshoppers that con-
sumed a single versus several plant species. In some trials,
a single plant was fed upon, and in others, all plants were
sampled. Grasshoppers have been shown to feed on one
plant during preference studies and then, on the following
day, show a signicantly different preference (Behmer and
Joern 1993; Howard 1993). In addition, grasshoppers are
known to sample many plant species prior to feeding upon
a single species (Haldar et al. 1995). In our experiments,
none of the A. xanthoptera individuals consumed all four
plant species, nor did they all avoid the same plant. The
fact that 28% of D. viridis fed on all four plants offered
suggests that they are either more generalist feeders or that
they sample many potential food plants prior to feeding.
Smooth brome and Kentucky bluegrass are sod-
forming grasses. These species have the capacity to form
near-monocultures and thus may be more “apparent” to
herbivores than more sparsely or patchily distributed spe-
cies, such as big bluestem and sideoats grama, which are
both bunchgrasses. Because plants that are more apparent
have a higher probability of insect attack, they are more
likely to employ constitutive (rather than induced) defenses
(Zangerl and Rutledge 1996). We would also expect the non-
native grasses we tested to be more heavily defended than
the native bunchgrasses tested. Defenses against herbivory
can allow some invasive plant species to outcompete native
plants, displacing them. Such a phenomenon is often ex-
plained in terms of “the enemy release hypothesis,” which
predicts exotic plants to become invasive when they leave
behind specialist herbivores from their native range and are
successful in deterring generalist herbivores in their new
range (Keane and Crawley 2002). In their study of feeding
by two native generalist grasshoppers on various invasive
and noninvasive plant species, Jogesh et al. (2008) found a
negative correlation between feeding and level of plant in-
vasiveness in one species of grasshopper but not the other.
While we did not directly study defensive compounds, our
results suggest that the autumn and green grasshopper are
not deterred from feeding on smooth brome and Kentucky
bluegrass, despite the expectation that these two species
would be well-defended against herbivory.
Grasshoppers can exert a strong effect on plant com-
munity composition. Branson and Sword (2008) found that
grasshopper herbivory reduced native plant species rich-
ness and abundance in a community dominated by crested
wheatgrass. We found that the two tested species preferred
non-native cool-season grasses over native grasses. Po-
tentially, these insect species can aid in maintaining plant
diversity in remnant prairies (Porter and Redak 1997) by
differentially feeding upon invasive grasses. Identifying
the roles grasshoppers may play in conserving native
prairie should become a research priority. The impacts of
grasshopper herbivory on plants that decrease or increase
under heavy grazing should also be investigated to better
The Grasshoppers Arphia xanthoptera and Dichromorpha viridis • Sean D. Whipple et al. 185
© 2009 Center for Great Plain s Studies , University of Ne braska–L incoln
understand the interplay between invertebrate herbivores
and grazing livestock in determining plant community char-
acteristics. Further study to identify feeding preferences for
individual grasshopper species would provide insights into
these ecological interactions and guide management deci-
sions that impact both plants and grasshoppers.
ACKNOWLEDGMENTS
The authors thank Drs. David Smith, John Hastings,
and Tamara Smith for useful discussion and comments
on earlier versions of this paper. We also thank three
anonymous reviewers for helpful suggestions to improve
this manuscript. In addition, we thank Rachel Anschutz
for assistance in collecting plants. This project was par-
tially supported by the University of Nebraska Research
Services Council.
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