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Received: 14 June 2018    Accepted: 12 December 2018

DOI: 10.1111/1365-2435.13272

RESEARCH ARTICLE

Fire, grazing and climate shape plant–grasshopper interactions

in a tallgrass prairie

Ellen A. R. Welti1  | Fan Qiu2 | Hannah M. Tetreault3 | Mark Ungerer2 | John Blair2 | 

Anthony Joern2

1
Department of Biology, The University of
Oklahoma, Norman, Oklahoma Abstract
2
Division of Biology, Kansas State University, 1. Species interactions are integral to ecological community function, and the struc‐
Manhattan, Kansas
ture of species interactions has repercussions for the consequences of species
3
USDA, ARS, University of Nebraska,
extinctions. Few studies have examined the role of environmental factors in con‐
Lincoln, Nebraska
trolling species interaction networks across time.
Correspondence
2. We examined variation in plant–grasshopper network structural properties in re‐
Ellen A. R. Welti
Email: welti@ou.edu sponse to three major grassland drivers: periodic fire, ungulate grazing and
climate.
Funding information
NSF DEB, Grant/Award Number: 1020465, 3. We sequenced a plant barcoding gene from extracted grasshopper gut contents
1342787 and 1440484
to characterize diets of 26 grasshopper species. Resulting grasshopper species’
Handling Editor: Rana El‐Sabaawi diets were combined with long‐term plant and grasshopper surveys to assemble
plant–grasshopper networks across 13–19 years for six watersheds subjected to
varying fire and grazing treatments.
4. Network modularity, generality and predicted grasshopper community robust‐
ness to plant species loss all increased in grazed watersheds. Temperature de‐
creased predicted grasshopper community robustness to plant species loss.
5. Grasshopper communities were found to be vulnerable to climatic warming due to
host plant loss. However, intermediate disturbance from ungulate grazers may
maintain grasshopper diversity and buffer community robustness to species loss.
Our results suggest that climate and disturbance shape the structure of ecological
interaction networks and thus have many indirect effects on species persistence
though direct effects on interaction partners.

KEYWORDS
Acrididae, community ecology, ecological networks, generalism, grasslands, herbivory,
modularity, robustness

1 |  I NTRO D U C TI O N
Plant–herbivore interactions are critical building blocks of terres‐
trial food webs, where plant species comprise ~20% and herbivo‐
rous insects ~25% of known non‐microbial terrestrial biodiversity
(Price, 2002). Ascertaining plant–herbivore network composition
is challenging as it requires identification of both species and diet
interactions (Novotny & Basset, 2005). Moreover, recent efforts
have only begun to examine how habitat and climatic conditions
shape network structural properties (Dalsgaard et al., 2013; López‐
Carretero, Díaz‐Castelazo, Boege, & Rico‐Gray, 2014; Rezende,
Albert, Fortuna, & Bascompte, 2009; Welti & Joern, 2015).
Structural properties of networks such as modularity and gen‐
erality of species interactions (diet breadth) provide insight into

Functional Ecology. 2019;1–11. © 2018 The Authors. Functional Ecology |  1

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© 2018 British Ecological Society
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2       WELTI et al.
ecological resilience. A modular species interaction structure results
when species interact primarily within sub‐modules or groups of
highly connected species (Montoya, Pimm, & Sole, 2006; Rezende
et al., 2009; Thébault & Fontaine, 2010). Modularity decreases ex‐
tinction risk by limiting the spread of perturbation due to species
loss outside of a specific subgroup. Modularity reduces competi‐
tion among species (Rezende et al., 2009) and decreases sensitivity
to fluctuation of inter‐module species (Montoya et al., 2006). The
degree of herbivore diet generalism is also predicted to buffer her‐
bivores against plant species loss. As generalist herbivore species
can use alternate plant hosts, they should be less susceptible to
fluctuations in resource availability (López‐Carretero et al., 2014).
Additionally, robustness to extinction of an ecological community
can be predicted through extinction simulation models (Memmott,
Waser, & Price, 2004). For example, use of these models can quan‐
tify the tolerance of the herbivore community to plant species loss
(Burgos et al., 2007).
Here, we examine changes in plant–grasshopper networks in a
Kansas tallgrass prairie, an ecosystem shaped by exposure to peri‐
odic fire, grazing and climate variability. In this system, fire and graz‐
ing regimes are known to modify grasshopper communities directly
(Jonas & Joern, 2007) and indirectly through their effects on plants
(Hartnett, Hickman, & Fischer‐Walter, 1996). Preferential grazing on
grasses by bison reduces the competitive pressures on forbs (Towne,
Hartnett, & Cochran, 2005), increases overall plant species diver‐
sity and increases grasshopper diversity in response to changes to
plant community composition (Joern & Laws, 2013). Increases in
fire frequency indirectly affect grasshopper communities by re‐
moving accumulated plant litter, altering the soil and near surface
environment, facilitating fire‐tolerant plant species and enhancing
dominance of several species of warm‐season grasses, thus modi‐
fying host plant availability for the grasshopper community by de‐
creasing available plant diversity. Fire events produce concentrated
but transient pulses of nutrients (e.g. P and cations) and long‐term
reductions in others (e.g. N) (Blair, 1997). Additionally, fire increases
primary production in the absence of grazing (Collins & Steinauer,
1998). Consequently, some grasshopper species are highly respon‐
sive to variation in historical fire frequency (Joern, 2005).
Long‐term data facilitate observational examination of the ef‐
fects of climate on ecological communities. Annual variation in pre‐
cipitation and temperature directly affects grasshopper physiology
and indirectly affects grasshopper populations through effects
on host plants, predators and pathogens (Jonas & Joern, 2007).
Increases in precipitation generally increase host plant quantity and
species richness, but can reduce plant nutritional quality (Cleland et
al., 2013; Mopper & Whitham, 1992). While the effects of drought
stress on host plants can increase grasshopper growth and repro‐
ductive success for individual species (Franzke & Reinhold, 2011),
more extreme drought can reduce grasshopper species richness
(Lenhart, Eubanks, & Behmer, 2015). Fungal pathogens can have
major impacts on grasshopper populations (Kistner & Belovsky,
2016), and these pathogens often increase in wetter years (Brust,
Hoback, & Wright, 2007). Temperature can likewise alter host plant
quality and quantity and directly affect grasshopper developmental
rates (Carter, Macrae, Logan, & Holtzer, 1998; Mukerji & Randell,
1975). In sum, the effects of climate on grasshopper populations and
communities are many, and understanding these dynamics requires
long‐term observation.
Here, we couple DNA barcoding of gut contents of 26 coexist‐
ing grasshopper (Acrididae) species with long‐term grasshopper and
plant datasets from a Kansas tallgrass prairie, to assemble putative
plant–grasshopper networks across 13–19 years. We calculated
plant–grasshopper interaction weights for each interaction for each
watershed and year using (a) proportion of plant species in grasshop‐
per species diets, (b) grasshopper species frequency and (c) mean
plant species cover. We then compared plant–grasshopper networks
across watersheds subjected to different fire and grazing treatments
to characterize effects of these grassland disturbances on plant–
grasshopper network structural properties.
We predicted that through enhanced habitat heterogeneity, nu‐
trient availability and plant species diversity, ungulate (bison) graz‐
ing would lead to shifts in plant–grasshopper interaction structure.
Predicted shifts in network structure due to the presence of grazers
include the following: (a) increased modularity, through increases in
plant species richness allowing the addition of plant–grasshopper
network modules, especially increases in forbs due to bison grazing
and thus increases in forb‐feeding grasshoppers (Hickman, Hartnett,
Cochran, & Owensby, 2004); (b) increased grasshopper feeding gen‐
erality due to increased plant species richness supporting greater
abundance of grasshopper species with a wider diet breadth
(Bernays, Bright, Gonzalez, & Angel, 1994; Unsicker et al., 2010);
and (c) increased grasshopper community robustness to local plant
species loss due to increased plant species redundancy in grasshop‐
per diets. Because of its negative effect on plant species diversity,
we expected increased fire frequency to decrease the modularity,
generality and robustness of plant–herbivore networks, especially in
the absence of grazers. Increases in annual precipitation could either
increase modularity, generality and robustness by increasing host
plant availability or decrease these network structural properties
by decreasing food quality or decreasing grasshopper abundance
due to negative effects of grasshopper fungal pathogens. Increase
in mean annual temperature was predicted to decrease modularity,
generality and robustness as higher temperatures stress both plant
and grasshopper communities.
2 | M ATE R I A L S A N D M E TH O DS
2.1 | Overview
To assess the influence of fire, grazing and climate on plant–grass‐
hopper network structure, we merged our snapshot dataset of
grasshopper diets with two long‐term datasets to assemble model
networks. Specifically, we used 13–19 years of plant and grasshop‐
per monitoring data collected in watersheds subjected to different
treatments of fire (three frequencies) and bison grazing (present,
excluded) to determine abundance and identify of participants in
WELTI et al.
plant–grasshopper networks. These datasets were combined with
grasshopper diet data, estimated from DNA barcoding, to identify
likely feeding interactions across time and space.
2.2 | Site description
Grasshopper and plant data were collected at Konza Prairie Biological
Station (KPBS), a 3,487 ha native tallgrass prairie in NE Kansas, USA.
KPBS had a mean annual precipitation of 835 mm and a mean annual
temperature of 12.9°C over the period of this study (1996–2014).
Beginning in 1977, a landscape‐level fire‐frequency experiment has
been conducted at KPBS, with individual watersheds subjected to
fire‐frequency treatments of 1, 2, 4 or 20 years. In 1987, bison were
reintroduced to a portion of KPBS. The current herd of ~280 ani‐
mals has free access to 10 watersheds (~1,000 ha), encompassing an
area containing watersheds subjected to replicated fire‐frequency
treatments.
2.3 | Konza LTER datasets
Long‐term grasshopper and plant community composition data‐
sets from selected KPBS watersheds were collected as part of the
National Science Foundation Long‐Term Ecological Research (NSF
LTER) programme. Datasets and detailed information on sampling
protocols are available on the KPBS LTER website: http://lter.
konza.ksu.edu/data-explorer-dashboard (grasshopper dataset code:
CGR02; plant dataset code: PVC02). Six watersheds included in both
datasets and consisting of a crossed experimental design of bison
grazing (three watersheds grazed by bison and three non‐grazed wa‐
tersheds) and fire frequency (two watersheds of each with fire re‐
turn intervals of 1, 4 and 20 years) were selected for analysis in this
study (watershed codes: 1D, 4B, 20B, N1B, N4D and N20B).
Data from 1996 to 2014 (19 years) were analysed for the three
non‐grazed watersheds. Grasshopper sampling on grazed water‐
sheds did not begin until 2002; data from 2002 to 2014 (13 years)
were analysed for the three grazed watersheds. Briefly, grasshop‐
pers for the KPBS long‐term grasshopper dataset were sampled
using sweep nets twice per year in midsummer at two locations per
watershed. At each location and date, ten samples were taken, each
sample consisting of 20 sweeps (40 samples/watershed/year and
800 sweeps/watershed/year). The frequency of each grasshopper
species was calculated as the proportion of the 40 samples in which
the grasshopper species was collected.
Long‐term plant composition at KPBS was sampled across eight
50 m transects/watershed, each consisting of five 10‐m2 plots. For
each plant species present in each plot, a per cent cover category
of 0%–1%, 2%–5%, 5%–25%, 25%–50%, 50%–75%, 75%–95% or
95%–100% was recorded. We used the median of each selected
range for each species in each plot (Ratajczak, Nippert, & Ocheltree,
2014) and averaged across plots within a watershed and year to es‐
timate mean per cent cover of each plant species/watershed/year.
For plant species identified from DNA extracted from grasshopper
Functional Ecology |
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gut content to only the genus level, all plant species within the genus
were averaged.
Precipitation and temperature data were collected from the KPBS
meteorological station and archived by the Climate and Hydrology
Database Project with support from the NSF LTER and USDA Forest
Service (http://climhy.lternet.edu/). A state‐level index of drought
conditions, the Palmer Drought Severity Index (PDO), was down‐
loaded from the National Climate Data Center climate monitoring
website (https://www.ncdc.noaa.gov/cag/time-series/us).
2.4 | Establishing grasshopper diets
Grasshoppers used for diet analyses were collected with sweep
nets as encountered in all watersheds across KPBS in August 2014,
aiming for a collection across a wide distribution of habitat types
within each species. We attempted to collect 20 individuals (10 adult
males and 10 adult females) of each grasshopper species present;
however, samples sizes were smaller for uncommon species (total
grasshopper individuals = 452; see Table S1, Supporting Information
for number of individuals sampled per species). Twenty‐nine species
of grasshopper were collected, and we were able to extract, amplify
and sequence gut content DNA from 26 species. With the exception
of one relatively common grass‐feeding grasshopper (Mermiria bivit‐
tata) from which we were unable to extract DNA, these 26 species
comprise the majority of the egg‐overwintering grasshopper species
at KPBS.
2.5 | DNA extraction, PCR and Illumina
MiSeq sequencing
To characterize grasshopper diets, plant DNA was extracted
from the dissected crop of each individual grasshopper using
the DNeasy Plant Mini Kit (Qiagen, Valencia, CA) according to
manufacturer's protocol, quantified with a NanoDrop ND2000
Spectrophotometer (Thermo Scientific, Wilmington, Delaware,
USA) and templates standardized to 2 ng/µl. PCR was performed
with primers targeting the ~600 bp chloroplast trnL intron (for‐
ward primer: CGAAATCGGTAGACGCTACG; reverse primer:
GGGGATAGAGGGACTTGAAC12bp_tag) (Avanesyan, 2014; Taberlet,
Gielly, Pautou, & Bouvet, 1991). Reverse primers were synthesized
with unique 12 bp 3′ tags (Table S1, Supporting Information) to fa‐
cilitate multiplex sequencing and identification of samples originat‐
ing from the 26 grasshopper species. Each sample was amplified in
a 25 μl reaction consisting of 1 μM of forward and reverse primers,
10 ng of template DNA, 200 μM of dNTPs, 1 μM of MgCl2, 0.5 units
of proofreading Phusion Green Hot Start II High‐Fidelity DNA
Polymerase (Thermo Scientific, Wilmington, Delaware, USA) and
5 μl of 5X Green HF PCR Buffer (Thermo Scientific). PCR thermal
cycler parameters followed Taberlet et al. (1991). Briefly, PCR condi‐
tions were 98°C for 5 min, followed by 35 cycles with denaturation
at 94°C for 1 min, annealing at 50°C for 30 s, extension at 72°C for
1 min, with final extension at 72°C for 10 min. Negative controls
 | Functional Ecology
4       WELTI et al.
were performed where template DNA was withheld and none pro‐
duced amplicons.
To clean up PCR amplicons and remove residual PCR primers,
we used ExoSAP‐IT (Thermo Scientific) followed by the Agencourt
AmPure SPRI system (1:1 ratio of bead solution to PCR volume).
Purified PCR products were visualized on a 1.5% agarose (w/v) gel to
ensure successful amplification. Amplicon DNA concentration from
each grasshopper species was measured and pooled (300 ng per
species). The amplicon libraries were paired‐end sequenced using
the MiSeq Personal Sequencing System 600‐Cycle Kit (Illumina, San
Diego) at the Kansas State University Integrated Genomics facility.
2.6 | Bioinformatics
Sequencing data were processed and assembled into contigs using
mothur (version 1.33.3; Schloss et al. 2009). Sequences were re‐
moved using mothur (version 1.33.3) based on the following criteria:
(a) containment of any ambiguous bases, (b) more than two mis‐
matches to the primers, (c) any mismatches to unique 12 bp 3′ tags,
(d) homopolymers longer than 8 bp, (e) were <250 bp in length, (f)
any containing chimeric properties as determined using the UCHIME
algorithm (Edgar, 2010) and (g) any without a minimum overlap of
50 bp. This yielded 1,067,005 sequences. Sequences were then
clustered as operational taxonomic units (OTUs) based on a 97%
similarity using nearest neighbour joining that conservatively assigns
sequences to OTUs. Raw sequence reads have been submitted to
NCBI Short Read Archive.
To identify the assembled sequences from each grasshopper spe‐
cies to plant taxa, the chloroplast trnL gene for all species belong to
plant genera found at KPBS (for list of KPBS plants, we used Towne,
2002) was downloaded from GenBank (31,765 sequences, defined as
the trnL database). Assembled sequences were compared to the trnL
database with the BLASTN tool of BLAST v2.2.31 (Altschul, Gish,
Miller, Myers, & Lipman, 1990). The best BLAST hit of each sequence
was used for plant identification (e‐value <10−20). Sequences from gut
contents matched 47 plants at the species level and 19 at the genus
level. Only plants with >5 sequence matches per grasshopper species
were used to assemble plant–grasshopper networks.
2.7 | Interaction weights
Interaction weights were calculated to assign a quantitative value
to the importance of each feeding interaction for a given grasshop‐
per species and plant species/watershed/year (modified from Welti,
Helzer, & Joern, 2017). Interaction weights for each plant species or
genus (i) and grasshopper species (j) interaction within each water‐
shed (w) and year (y) were calculated as follows:
Sij
Interaction weightijwy = ( ) × Piwy × Gjwy
Tj
where S is the number of sequences of the plant taxa (i) se‐
quenced from the grasshopper species (j)’s gut contents, T is the
total number of sequences from that grasshopper species (j)’s gut
contents (S/T is the estimated proportion of a grasshopper species
(j)’s diet comprised by a given plant (i)), P is the mean per cent plant
cover of plant taxa (i) from plant sampling within each watershed (w)
and year (y), and G is the frequency of sampled grasshopper species
(j) within each watershed (w) and year (y). Interaction weights can
range from 0 to 100 units of strength.
2.8 | Network analyses
Plant–grasshopper networks built from species composition and
calculated interaction weights were assembled for each water‐
shed and year (19 years*3 non‐grazed watersheds + 13 years * 3
grazed watersheds = 96 total networks). Network indices of gen‐
erality, modularity and robustness were calculated for each plant–
grasshopper network using the Bipartite package (Dormann,
Fründ, Blüthgen, & Gruber, 2009) in program R (version 3.3.0; R
Development Core Team, 2016). Generality (generality.HL, net‐
worklevel function) measures the weighted mean number of plant
species per grasshopper species (Bersier, Banasek‐Richter, &
Cattin, 2002, Tylianakis, Tscharntke, & Lewis, 2007). Modularity
(modularity, computeModules) measures the weighted degree
of compartmentalization using Newman (2004)’s algorithm
(Dormann & Strauß, 2013). Grasshopper robustness to plant spe‐
cies loss (robustness.HL, robustness function) measures the aver‐
age area under the curve of 1,000 simulations of the relationship
between random removal of plant species (primary extinctions)
and predicted loss of grasshopper species (secondary extinctions)
(Burgos et al., 2007; Memmott et al., 2004). Plant–grasshopper
networks, network metrics and habitat data are archived in the
Dryad Digital Repository (https://doi.org/10.5061/dryad.s6j1822;
Welti et al., 2018).
2.9 | Comparison with previous grasshopper
diet studies
To evaluate the completeness of our diet dataset, we include here a
comparison with two previous grasshopper diets studies conducted
at sites in close proximity to our study location. Both of these studies
(Mulkern et al., 1969 and Campbell, Arnett, Lambley, Jantz, & Knutson,
1974) collected grasshoppers from a former native pasture managed
by the Agricultural Experimental Station of Kansas State University,
within 15 km of KPBS, and identified plant fragments from dissected
grasshopper guts under a microscope. Using these data, we calculated
grasshopper diet generality, modularity, grasshopper robustness and
network connectance to compare with our data. All network metrics
were calculated using the Bipartite package (Dormann et al. 2009) in
program R (version 3.3.0; R Development Core Team, 2016).
2.10 | Statistical analyses
Global linear mixed models fit by REML for each of three network
indices (generality, modularity and robustness) included the fixed
WELTI et al.
predictor variables of grazing (presence or absence), fire frequency
(1, 4, or 20 year burn intervals), years since last fire, annual cumula‐
tive precipitation, mean annual temperature and the Palmer Drought
Severity Index. Individual watersheds were included as a random
variable in models to account for repeated observations (Zuur, Ieno,
Walker, Saveliev, & Graham, 2009). Linear mixed models were fitted
using the lme4 package (Bates, Maechler, Bolker, & Walker, 2015),
and conditional R2 values were calculated following Nakagwa and
Schielzeth (2013) in program R (version 3.3.0; R Development Core
Team, 2016). Model residuals were regressed over year to assess
temporal autocorrelation, and no significant correlations were found
(Figure S1, Supporting Information). As watersheds included in this
study are not replicated at the level of combined fire and grazing
treatments, we are not able to statistically test for fire and grazing
interactions.
Models predicting plant–grasshopper network structural prop‐
erties using fire and grazing treatments and climate variables were
compared using Akaike information criteria (AIC) (Burnham &
Anderson, 2002). Global (models including all predictor variables)
and all reduced models (all non‐interactive combinations of predic‐
tors) were compared using the dredge function (MuMIn package,
[Bartoń, 2016]) in program R (version 3.3.0, R Development Core
Team, 2016). Models were considered competitive and equally
parsimonious when they had a ∆AICc < 2 (Burnham & Anderson,
2002). Relative importance values were calculated as the sum of all
AIC weights containing a predictor variable (Burnham & Anderson,
2002). As AIC results can be difficult to visualize, we use regression
plots to depict changes in network structure.
A partial redundancy analysis (RDA) ordination was used to ex‐
amine how fire and grazing shaped plant and grasshopper commu‐
nities (Borcard, Legendre, & Drapeau, 1992; Legendre & Anderson,
1999). Additionally, RDAs relating plant and grasshopper community
composition and modularity, grasshopper generality and robust‐
ness were used to determine whether community composition im‐
pacted plant–grasshopper network structure. RDAs were conducted
using the vegan package (Oksanen et al., 2015) in R ver. 3.3.3 (R
Development Core Team, 2016).
3 | R E S U LT S
Grasshopper gut content analyses revealed 572 individual feed‐
ing interactions between 26 grasshopper species and 66 plant taxa
(species or genera). The full plant–grasshopper interaction web is
shown in Figure S2, Supporting Information. Plant–grasshopper
communities in grazed watersheds had more species and interac‐
tions than those in non‐grazed watersheds (Figure 1), and the pres‐
ence of grazers increased modularity, generality and grasshopper
robustness (Figure 2; Figure S3, Supporting Information). Bison
grazing and mean annual temperature are top drivers of plant–grass‐
hopper network structure (Figure 3). Fire frequency and grazer
treatments had significant effects on plant and grasshopper com‐
munity composition (Figure S4, Supporting Information). Plant and
Functional Ecology |
      5
grasshopper community composition was significantly correlated
with plant–grasshopper network structure (Figure S5, Tables S2 and
S3, Supporting Information).
3.1 | Modularity
Grazer treatment and mean annual temperature had the highest
relative importance values (Figure 3), and grazer treatment was
included in the top AIC c predicting modularity of plant–grass‐
hopper networks (Table S4, Supporting Information). Modularity
was higher in watersheds grazed by bison (Figure 2, Panel a and
d; Figure S3, Panel A, Supporting Information). Within the non‐
grazed watersheds, modularity of plant–grasshopper networks
was highest in the 20‐year burn frequency (Figure S3, Panel A,
Supporting Information). Modularity tended to decrease with
mean annual temperature, but this trend was highly variable and
not significant (F1,94 = 3.3, R 2 = 0.02, p = 0.07; Figure 2, Panel d).
Across all watersheds, cumulative annual precipitation did not
have a significant linear effect on modularity (F1,94 = 0.4, R 2 < 0.01,
p = 0.51; Figure 2, Panel a).
3.2 | Generality
Bison grazing had the largest effect of grassland drivers on generality
(Figure 3) where grazing was the only fixed effect appearing in top AICc
models (Table S4, Supporting Information). While the null model (model
including only watershed to account for repeated observations) was
also included in top models, grazer presence increased grasshopper
diet generality between watersheds (Figure 2, Panels b and e; Figure
S3, Panel B, Supporting Information). Within grazed watersheds, gen‐
erality was higher and more variable in 4‐year burns (Figure 2, Panels
b and e). Although present in top models, relationships between gen‐
erality and mean annual temperature (F1,94 = 2.0, R2 = 0.01, p = 0.16;
Figure 3, Panel e) or cumulative annual precipitation (F1,94 = 1.3,
R2 < 0.01, p = 0.25; Figure 3, Panel b) were not significant.
3.3 | Grasshopper robustness
Mean annual temperature was the top driver (Figure 3, Table
S4, Supporting Information), decreasing grasshopper robustness
(F1,94 = 22.6, R2 = 0.19, p < 0.001; Figure 3, Panel f). Additionally,
grasshopper robustness was higher in grazed watersheds (Figure 2
Panels c and f; Figure S3, Panel C, Supporting Information). Fire‐fre‐
quency effects were not significant, although watersheds burned
every 4 years tended to have higher grasshopper robustness
(Figure 2, Panels c and f). Grasshopper robustness increased with
cumulative annual precipitation (F1,94 = 15.5, R2 = 0.13, p < 0.001;
Figure 3, Panel c). The driest (569 mm cumulative precipitation) and
hottest (mean 14.9°C) year (2012) had low plant species richness,
driving climatic relationships and suggesting a threshold for grass‐
hopper robustness. While temperature and precipitation had inverse
correlations with robustness, these two climate variables were not
correlated across 1996–2014 (F1,17 = 1.3, R2 = 0.02, p = 0.27).
 | Functional Ecology
6       WELTI et al.

F I G U R E 1   Comparison of representative plant–grasshopper networks from the most recent year of data (2014). The top yellow bars
represent grasshopper species, and the bottom green bars represent plant species in the network. The blue lines connecting plant and
grasshopper species represent feeding links, and the thickness of the lines corresponds to the strength of the species’ interaction. Network
panels are arranged by treatment with row indicating grazing treatment and columns indicating fire return interval. The annually burned and
not grazed watershed in the top left is dominated by two grasshopper species (Orphulella speciosa and Phoetaliotes nebrascensis) and one
plant species (Andropogon gerardii), whereas other fire and grazing treatments resulted in more complex grasshopper–plant networks

F I G U R E 2   Relationships between climate drivers of cumulative annual precipitation (a–c) and mean annual temperature (d–f) and plant–
grasshopper network structures of modularity, generality and robustness to plant species loss

1969) and Figure S7, Supporting Information (Campbell et al., 1974).

3.4 | Comparison with other plant–grasshopper
diet studies
Plant–grasshopper networks from previous grasshopper diet studies
are depicted in Figure S6, Supporting Information (Mulkern et al.,
Compared to our 26 grasshopper and 66 plant species, the Mulkern
et al. (1969) study included 19 grasshopper and 47 plant species
and the Campbell et al. (1974) study included 17 grasshopper and
58 plant species. With the possible exception of lower grasshopper
WELTI et al.
1 Grazing
0.9
Relave importance value
0.8
0.64
0.7 0.63
0.6
0.5
0.4
0.24
0.3
0.2
0.1
0.01
0.01
0.01
0.01
0 levels of modularity and generality occurred in watersheds burned
Modularity
F I G U R E 3   Relative importance values, a summed and
standardized indicator of predictor variable rank across all possible
models, for grassland driver predictors (grazing = presence/absence
of bison grazers, fire = historical burn interval, YSB = years since
last burn, CAP = cumulative annual precipitation, MAT = mean
annual temperature, PDI = Palmer drought index) of the network
structural properties of grasshopper robustness, modularity and
generality of grasshopper diets. Top models with ∆AICc < 2 are
given in Supporting Information Table S4
generality found in this study, all plant–grasshopper diet studies had
similar network structure values. The lower grasshopper generality
in our study likely reflects differences in environments; Mulkern et
al. (1969) and Campbell et al. (1974) were both performed in solely
grazed pastures with high stocking densities. Additionally, our study
includes more grasshopper species, potentially capturing more spe‐
cialist species.
4 | D I S CU S S I O N
While insect communities in particular can be influenced by a suite
of environmental factors (Andrewartha & Birch, 1954; Farkas &
Singer, 2013), identifying which factors are most important remains
a major challenge, and long‐term studies are needed to tease apart
driver contributions to community composition and dynamics (Jonas
& Joern, 2007; Lenhart et al., 2015; Olesen, Bascompte, Elberling,
& Jordano, 2008) and interaction patterns (Thébault & Fontaine,
2010). Using long‐term plant and grasshopper community datasets,
we found grazing and mean annual temperature shaped grasshop‐
per–plant network structure. Bison grazing increased modularity
and generality, whereas mean annual temperature decreased grass‐
hopper robustness.
Grazing impacted the plant and grasshopper communities, in‐
creasing species richness in grazed watersheds and altering commu‐
nity composition. Large ungulates directly modify plant community
composition and vegetation structure through preferential grazing
(Hartnett et al., 1996; Jonas & Joern, 2007). While it is counter‐in‐
tuitive that modularity and generality should increase together, this
grazing response is due to the increase in a large module of gen‐
eralist forb‐feeding grasshoppers. Previous studies documented
both positive (Joern, 2005; Jonas & Joern, 2007; Zhong et al., 2014)
and negative (Belovsky, 1984; Branson & Haferkamp, 2014) grazing
Functional Ecology |
      7
effects on grasshopper communities. Here, bison grazing increased
1
Burning plant–grasshopper network structural properties of modularity, gen‐
erality and grasshopper robustness, suggesting grazing increases
YSB
CAP grasshopper community resilience to loss of host plant species.
MAT Changes in fire frequency affected plant and grasshopper com‐
PDI munity composition but did not have significant main effects on
plant–grasshopper network structure. We do not have replicated
0.22
0.21
watersheds at the level of fire and grazing treatments to test for a
0.07
fire‐grazing interaction. However, in grazed watersheds, the highest
0.04
0.01
0.01
0.01
0.01
0.01
0.01
Generality Robustness
every four years. This intermediate fire frequency combined with
grazing may therefore optimize plant–grasshopper network struc‐
tural properties.
Climate change predictions for central North American grass‐
lands generally include warmer temperatures and increased pre‐
cipitation variability (Dobrowski et al., 2013; Kunkel et al., 2013).
Temperature decreased grasshopper robustness to plant species
loss, likely mediated through shifts in plant community composition.
The dominant grass species at KPBS, Andropogon gerardii, decreases
in biomass at soil temperatures >25°C (DeLucia, Heckathorn, & Day,
1992). Based on previously reported plant community responses
(Harrison, Gornish, & Copeland, 2015) and our results, increasing
temperatures decrease plant species richness in this system, mak‐
ing grasshopper communities more susceptible to additional plant
loss (Schleuning et al., 2016) and decreasing grasshopper diversity
(Lenhart et al., 2015). High temperatures may also directly reduce
grasshopper populations if species pass thermal limits; a previous
temperature manipulation study on KPBS showed grasshopper sur‐
vival in the absence of predators was highest in the lowest tempera‐
ture treatment (Laws & Joern, 2012).
The lack of a significant precipitation or drought response in
our AICc model could reflect our reduced power to detect climatic
effects compared to fire and grazing effects, as fire and grazing
treatments are replicated across watersheds and the treatment dif‐
ferences may become stronger over time. The extreme reduction
in grasshopper robustness across all watersheds in 2012 suggests
a potential threshold for climatic drivers of plant–grasshopper net‐
work structure. Capturing changes in years of climatic extremes is an
advantage of long‐term data collection.
Modularity and generalism of herbivore diet are hypothesized
to be important structural property stabilizing bipartite plant–her‐
bivore networks (Montoya et al., 2006; Thébault & Fontaine, 2010).
While few previous studies have compared trophic networks within
the same system, comparisons across datasets have shown an in‐
crease in generality with species richness across producer–consumer
network comparisons, whereas the relationship between modular‐
ity and species richness is less clear (Morris, Gripenberg, Lewis, &
Roslin, 2014; Welti & Joern, 2015). However, while larger ecological
networks themselves may be more stable, modularity, generality and
robustness provide predictions for responses to future species loss.
Here, the more generalist and modular plant–grasshopper networks
in the grazed and intermediately burned watersheds are predicted to
be the most resilient to future plant species loss.
 | Functional Ecology
8       WELTI et al.
4.1 | Caveats
Our analysis of plant–grasshopper interaction networks was based
on quantification of grasshopper diets coupled with long‐term plant
and grasshopper community composition datasets, which provided
a unique opportunity to evaluate changes in plant–grasshopper in‐
teractions across years. We acknowledge several key assumptions
of these analyses. While only mean annual temperature, cumulative
annual precipitation and PDO were examined as explanatory driv‐
ers of plant–grasshopper networks, other measures such as climate
variability and timing of precipitation are likely important divers of
plant and grasshopper communities. We quantified grasshopper diet
composition using grasshoppers collected in 2014 and then extrapo‐
lated results to examine plant–grasshopper networks over multiple
years. We cannot assess intraspecific diet variation with our dataset
as we used unique DNA barcodes for each grasshopper species but
not each grasshopper individual. We assume that diet preferences
of grasshopper species do not change substantially among years
and that our constructed annual plant–grasshopper diet matrices,
based on annual changes in plant species availability and grasshop‐
per species population levels, are a reasonable representation of
grasshopper diets across years. Moreover, using the number of DNA
sequences to estimate proportion of diet can lead to potential error
in bias for or against particular plant species during grasshopper di‐
gestion, DNA extraction, amplification and sequencing (Pompanon
et al., 2012). However, bias exists in all methods of diet sampling.
Identification of plant fragments from dissected gut contents is bi‐
ased towards plants with tough cuticles (Leslie, Vavra, Starkey, &
Slater, 1983), and DNA barcoding has been shown to provide better
taxonomic resolution (Soininen et al., 2009). The relative number of
sequences represents our best estimates of proportion of plant spe‐
cies in a diet from this study, and its influence on interaction weight
is tempered by inclusion of grasshopper species frequency and plant
species cover in interaction weight calculation.
While we may have missed rare species, we do not expect major
annual shifts in grasshopper diet in terms of plant species identity
(Joern, 1979). Grasshopper species included in analyses but under‐
sampled may be misrepresented as more specialist; we opted to
include them in analyses to include all available information; how‐
ever, as most are rarely collected in long‐term annual monitoring,
their impact on network structure is minimal. Networks from pre‐
vious studies from a nearby location found similar network struc‐
tures for the full diet matrix (Figure S8, Supporting Information).
Insect herbivore diets are often conserved (Bernays & Bright, 1993;
Pérez‐Harguindeguy et al., 2003; Rowell‐Rahier, 1984); thus, we do
not expect species‐specific diets to shift significantly among years.
Additionally, grasshoppers are presumed to have similar diets across
life stages (Cárdenas, Gallardo, Moyano, & Presa, 2017). While in‐
sect herbivores are known to adjust diet selection in response to
plant species abundances (Bernays & Chapman, 1970; Huang,
McNeill, & Zhang, 2016; Singer & Stireman, 2001; Ueckert, Hansen,
& Terwilliger, 1972), we specifically account for variation resulting
from shifts in plant species abundances and grasshopper species
abundances, as mean plant cover and grasshopper frequency are
included in our calculation of interaction weights. Combining these
datasets allowed us to create a novel dataset of long‐term plant–
grasshopper interactions and thus examine possible major habitat
and climatic effects on plant–grasshopper interaction structures.
5 | CO N C LU S I O N S
Ungulate grazing and possibly intermediate fire frequency were
found to increase the modularity and generality of grasshopper diets
and increase grasshopper robustness to plant species loss. Grazing
by cattle, the functional replacement for native bison in the majority
of the remaining intact grasslands of North America, may maintain
grasshopper community resilience at moderate stocking densities.
However, homogenization of rangeland grassland systems through
annual burning and high stocking densities (Gossner et al., 2016)
is likely to reduce stabilizing structures of plant–grasshopper net‐
works. Only through long‐term monitoring can we examine antici‐
pated effects of climate trends on ecological communities. Although
it is difficult to predict community robustness to directional cli‐
matic shifts, our results for plant–grasshopper communities studied
here predict that anticipated warmer and drier conditions in North
American grasslands will decrease grasshopper robustness to plant
species loss, making them more vulnerable to local extinction.
AC K N OW L E D G E M E N T S
We are grateful to Allison Veach for her assistance with Bioinformatics.
Alina Akunova cleaned PCR amplicons and conducted Illumina se‐
quencing, and Lydia Zeglin supplied laboratory space for conducting
DNA clean‐up steps. Thanks to Keith Gido and Gregory Zolnerowich
for their feedback which substantially improved this manuscript.
We thank Konza Prairie LTER for data access. Grants from NSF DEB
1020465, NSF DEB 1342787, NSF DEB 1440484 and Konza Prairie
Biological Station supported this research. E.A.R.W. was supported
by a Graduate Assistance in Areas of National Need fellowship from
the U.S. Department of Education.
AU T H O R S ' C O N T R I B U T I O N S
E.A.R.W. and A.J. designed the study. E.A.R.W. collected and ana‐
lysed data and wrote the first draft of the manuscript. H.M.T. as‐
sisted with DNA extraction and PCR. F.Q. conducted bioinformatics.
M.U. and J.B. proved technical support and conceptual advice. All
authors discussed results and substantially contributed to manu‐
script revisions.
DATA AC C E S S I B I L I T Y
Plant–grasshopper networks, network metrics and habitat data de‐
posited in the Dryad Digital Repository: https://doi.org/10.5061/
dryad.s6j1822 Welti et al., 2018. Raw sequence reads are
WELTI et al.
available from the NCBI Short Read Archive (SRA BioProject ID
PRJNA509933).
ORCID
Ellen A. R. Welti  https://orcid.org/0000-0001-6944-3422
REFERENCES
Altschul, S. F., Gish, W., Miller, W., Myers, E. W., & Lipman, D. J. (1990).
Basic local alignment search tool. Journal of Molecular Biology, 215,
403–410. https://doi.org/10.1016/S0022-2836(05)80360-2
Andrewartha, H. G., & Birch, L. C. (1954). The distribution and abundance
of animals. Chicago, IL: University of Chicago Press.
Avanesyan, A. (2014). Plant DNA detection from grasshopper guts: A
step‐by‐step protocol, from tissue preparation to obtaining plant
DNA sequences. Applications in Plant Sciences, 2, 1300082. https://
doi.org/10.3732/apps.1300082
Bartoń, K. (2016). Package ‘MuMIn’. Model selection and model averaging
based on information criteria (AICc and alike). R package version 3.2.4.
Retrieved from https://cran.r-project.org/web/packages/MuMIn/
index.html
Bates, D., Maechler, M., Bolker, B., & Walker, V. (2015). Fitting linear
mixed‐effects models using lme4. Journal of Statistical Software, 67,
1–48.
Belovsky, G. E. (1984). Moose and snowshoe hare competition and a
mechanistic explanation from foraging theory. Oecologia, 6, 150–159.
https://doi.org/10.1007/BF00396753
Bernays, E. A., & Bright, K. L. (1993). Mechanisms of dietary mixing in
grasshoppers: A review. Comparative Biochemistry and Physiology,
104, 125–131. https://doi.org/10.1016/0300-9629(93)90019-Z
Bernays, E. A., Bright, K. L., Gonzalez, N., & Angel, J. (1994). Dietary mix‐
ing in a generalist herbivore – tests of two hypotheses. Ecology, 75,
1997–2006. https://doi.org/10.2307/1941604
Bernays, E. A., & Chapman, R. F. (1970). Food selection by Chorthippus
parallelus (Zetterstedt) (Orthoptera: Acrididae) in the field. Journal of
Animal Ecology, 39, 383–394. https://doi.org/10.2307/2977
Bersier, L. F., Banasek‐Richter, C., & Cattin, M. F. (2002).
Quantitative descriptors of food‐web matrices. Ecology, 83,
2394–2407. https://doi.org/10.1890/0012-9658(2002)083[239
4:QDOFWM]2.0.CO;2
Blair, J. M. (1997). Fire, N availability, and plant response in grasslands: A test
of the transient maxima hypothesis. Ecology, 78, 2359–2368. https://
doi.org/10.1890/0012-9658(1997)078[2359:FNAAPR]2.0.CO;2
Borcard, D., Legendre, P., & Drapeau, P. (1992). Partialling out the spatial
component of ecological variation. Ecology, 84, 511–525. https://doi.
org/10.2307/1940179
Branson, D. H., & Haferkamp, M. (2014). Insect herbivory and vertebrate
grazing impact food limitation and grasshopper populations during
a severe outbreak. Ecological Entomology, 39, 371–381. https://doi.
org/10.1111/een.12114
Brust, M. L., Hoback, W. W., & Wright, R. J. (2007). Immersion toler‐
ance in rangeland grasshoppers (Orthoptera: Acrididae). Journal of
Orthoptera Research, 16, 135–138. https://doi.org/10.1665/1082-64
67(2007)16[135:ITIRGO]2.0.CO;2
Burgos, E., Ceva, H., Perazzo, R. P. J., Devoto, M., Medan, D., Zimmermann,
M., & Maria Delbue, A. (2007). Why nestedness in mutualistic net‐
works? Journal of Theoretical Biology, 249, 307–313.
Burnham, K. P., & Anderson, D. R. (2002). Model selection and multimodel
inference: A practical information‐theoretical approach. New York, NY:
Springer‐Verlag.
Campbell, J. B., Arnett, W. H., Lambley, J. D., Jantz, O. K., & Knutson,
H. (1974). Grasshoppers (Acrididae) of the Flint Hills native tallgrass
Functional Ecology |
      9
prairie in Kansas. Agricultural Experimental Station, Kansas State
University, Manhattan, KS, 19, 72–145.
Cárdenas, A. M., Gallardo, P., Moyano, L., & Presa, J. J. (2017).
Autecology, feeding preferences and reproductive biology of
Chortippus (Glyptobothrus) vagans (Eversmann, 1848) (Orthoptera:
Gomphocerinae) in Mediterranean ecosystems. Bulletin of
Entomological Research, 107, 21–31.
Carter, M. R., Macrae, I. V., Logan, J. A., & Holtzer, T. O. (1998).
Population model for Melanoplus sanguinipes (Orthoptera: Acrididae)
and an analysis of grasshopper population fluctuations in Colorado.
Environmental Entomology, 27, 892–901.
Cleland, E. E., Collins, S. L., Dickson, T. L., Farrer, E. C., Gross, K. L.,
Gherardi, L. A., … Turnbull, L. (2013). Sensitivity of grassland plant
community composition to spatial vs. temporal variation in precip‐
itation. Ecology, 94, 1687–1696. https://doi.org/10.1890/12-1006.1
Collins, S. L., & Steinauer, E. M. (1998). Disturbance, diversity and species
interactions in tallgrass prairie. In A. K. Knapp, J. M. Briggs, D. C.
Hartnett, & S. L. Collins (Eds.), Grassland dynamics: Long‐term ecolog‐
ical research in tallgrass prairie (pp. 140–156). New York, NY: Oxford
University Press.
Dalsgaard, B., Trøjelsgaard, K., Martín‐González, A. M., Nogués‐
Bravo, D., Ollerton, J., Petanidou, T., … Olesen, J. M. (2013).
Historical climate‐change influences modularity and nestedness
of pollination networks. Ecography, 36, 001–010. https://doi.
org/10.1111/j.1600-0587.2013.00201.x
DeLucia, E. H., Heckathorn, S. A., & Day, T. A. (1992). Effects of soil tem‐
perature on growth, biomass allocation and resource acquisition of
Andropogon gerardii Vitman. New Phytologist, 120, 543–549. https://
doi.org/10.1111/j.1469-8137.1992.tb01804.x
Dobrowski, S. Z., Abatzoglou, J., Swanson, A. K., Greenberg, J. A.,
Mynsberge, A. R., Holden, Z. A., & Schwartz, M. K. (2013). The cli‐
mate velocity of the contiguous United States during the 20th cen‐
tury. Global Change Biology, 19, 241–251. https://doi.org/10.1111/
gcb.12026
Dormann, C. F., Fründ, J., Blüthgen, N., & Gruber, B. (2009). Indices, graphs
and null models: analyzing bipartite ecological networks. Open Ecology
Journal, 2, 7–24. https://doi.org/ 10.2174/1874213000902010007
Dormann, C. F., & Strauß, R. (2013). Detecting modules in quantitative
bipartite networks: The QuaBiMo algorithm. arXiv [q‐bio.QM] 1304,
3218.
Edgar, R. C. (2010). Search and clustering orders of magnitude faster than
BLAST. Bioinformatics, 26, 2460–2461. https://doi.org/10.1093/
bioinformatics/btq461
Farkas, T. E., & Singer, M. (2013). Can caterpillar density or host‐plant
quality explain host‐plant related parasitism of a generalist forest
caterpillar assemblage? Oecologia, 173, 971–983.
Franzke, A., & Reinhold, K. (2011). Stressing food plants by altering water
availability affects grasshopper performance. Ecosphere, 2, 1–13.
https://doi.org/10.1890/ES11-00095.1
Gossner, M. M., Lewinsohn, T. M., Kahl, T., Grassein, F., Boch, S., Prati,
D., … Allan, E. (2016). Land‐use intensification causes multitrophic
homogenization of grassland communities. Nature, 540, 266–269.
https://doi.org/10.1038/nature20575
Harrison, S. P., Gornish, E., & Copeland, S. (2015). Climate‐driven di‐
versity loss in a grassland community. Proceedings of the National
Academy of Sciences of United States of America, 112, 8672–8677.
https://doi.org/10.1073/pnas.1502074112
Hartnett, D. C., Hickman, K. R., & Fischer‐Walter, L. E. (1996). Effects of
bison grazing, fire and topography on floristic diversity in tallgrass
prairie. Journal of Rangeland Management, 49, 413–420. https://doi.
org/10.2307/4002922
Hickman, K. R., Hartnett, D. C., Cochran, R. C., & Owensby, C. E. (2004).
Grazing management effects on plant species diversity in tall‐
grass prairie. Journal of Range Management, 57, 58–65. https://doi.
org/10.2307/4003955
 | Functional Ecology
10       WELTI et al.
Huang, X., McNeill, M., & Zhang, Z. (2016). Quantitative analysis of
plant consumption and preference by Oedaleus asiaticus (Acrididae:
Oedipodinae) in changed plant communities consisting of three grass
species. Environmental Entomology, 45, 163–170.
Joern, A. (1979). Feeding patterns in grasshoppers (Orthoptera:
Acrididae): Factors influencing diet specialization. Oecologia, 38,
325–347. https://doi.org/10.1007/BF00345192
Joern, A. (2005). Disturbance by fire and bison grazing modulate grass‐
hopper assemblages in tallgrass prairie. Ecology, 86, 861–873.
Joern, A., & Laws, A. (2013). Ecological mechanisms underlying arthro‐
pod species diversity in grasslands. Annual Review of Entomology, 58,
19–36. https://doi.org/10.1146/annurev-ento-120811-153540
Jonas, J., & Joern, A. (2007). Grasshopper (Orthoptera: Acrididae)
communities respond to fire, bison grazing and weather in North
American tallgrass prairie: A long‐term study. Oecologia, 153, 699–
711. https://doi.org/10.1007/s00442-007-0761-8
Kistner, E. J., & Belovsky, G. E. (2016). Host dynamics determine re‐
sponses to disease: Additive vs. compensatory mortality in a grass‐
hopper–pathogen system. Ecology, 95, 2579–2588.
Kunkel, K. E., Stevens, L. E., Stevens, S. E., Sun, L., Janssen, E., Wuebbles,
D., … Dobson, J. G. (2013). Regional climate trends and scenarios
for the U.S. National Climate Assessment. Part 4. Climate of the U.S.
Great Plains NOAA Technical Report NESMDIS. Washington, DC: U.S.
Department of Commerce.
Laws, A. N., & Joern, A. (2012). Predator‐prey interactions in a grass‐
land food chain vary with temperature and food quality. Oikos, 122,
977–986. https://doi.org/10.1111/j.1600-0706.2012.20419.x
Legendre, P., & Anderson, M. J. (1999). Distance‐based redundancy
analysis: Testing multispecies responses in multifactorial eco‐
logical experiments. Ecological Monographs, 69, 1–24. https://doi.
org/10.1890/0012-9615(1999)069[0001:DBRATM]2.0.CO;2
Lenhart, P. A., Eubanks, M. D., & Behmer, S. (2015). Water stress in grass‐
lands: Dynamic responses of plants and insect herbivores. Oikos, 124,
381–390. https://doi.org/10.1111/oik.01370
Leslie, D. M. Jr, Vavra, M., Starkey, E. E., & Slater, R. C. (1983). Correcting
for differential digestibility in microhistological analyses involving
common coastal forages of the Pacific Northwest. Journal of Range
Management, 36, 730–732. https://doi.org/10.2307/3898197
López‐Carretero, A., Díaz‐Castelazo, C., Boege, K., & Rico‐Gray, V.
(2014). Evaluating the spatio‐temporal factors that structure net‐
work parameters of plant‐herbivore interactions. PLoS ONE, 9,
e110430. https://doi.org/10.1371/journal.pone.0110430
Memmott, J., Waser, N., & Price, M. (2004). Tolerance of pollination
networks to species extinctions. Proceeding of the Royal Society
of Biological Sciences, 271, 2605–2611. https://doi.org/10.1098/
rspb.2004.2909
Montoya, J., Pimm, S., & Sole, R. (2006). Ecological networks and their fra‐
gility. Nature, 442, 259–264. https://doi.org/10.1038/nature04927
Mopper, S., & Whitham, T. G. (1992). The plant stress paradox: Effects
on the pinyon sayfly sex ratios and fecundity. Ecology, 73, 515–525.
Morris, R. J., Gripenberg, S., Lewis, O. T., & Roslin, T. (2014). Antagonistic
interaction networks are structures independently of latitude and
host guild. Ecology Letters, 17, 340–349.
Mukerji, M. K., & Randell, R. L. (1975). Estimation of embryonic devel‐
opment in populations of Melanoplus sanguinipes (Fabr.) (Orthoptera:
Acrididae) in the fall. Acrida, 4, 9–18.
Mulkern, G. B., Pruess, K. P., Knutson, H., Hagen, A. F., Campbell, J. B., &
Lambley, J. D. (1969). Food habits and preferences of grassland grass‐
hoppers of the North Central Great Plains. Bulletin No. 481 of the
North Dakota Agricultural Experimental Station, Fargo, ND, USA.
Nakagawa, S., & Schielzeth, H. (2013). A general and simple method for
obtaining R 2 from generalized linear mixed‐effects models. Methods
in Ecology and Evolution, 4, 133–142.
Newman, M. E. J. (2004). Analysis of weighted networks. Physical Review
E, 70, 056131. https://doi.org/10.1103/PhysRevE.70.056131
Novotny, V., & Basset, Y. (2005). Host specificity of insect herbivores in
tropical forests. Proceedings of the Royal Society B, 272, 1083–1090.
Oksanen, J., Blanchet, F. G., Kindt, R., Legendre, P., Minchin, P. R., O'Hara,
R. B., … Wagner, H. (2015). vegan: Community Ecology Package. R pack‐
age version 2.2‐1. http://CRAN.R-project.org/package=vegan.
Olesen, J. M., Bascompte, J., Elberling, H., & Jordano, P. (2008). Temporal
dynamics in a pollination network. Ecology, 89, 1573–1582. https://
doi.org/10.1890/07-0451.1
Pérez‐Harguindeguy, N., Díaz, S., Vendramini, F., Cornelissen, J. H. C.,
Gurvich, D. E., & Cabido, M. (2003). Leaf traits and herbivore se‐
lection in the field and in cafeteria experiments. Austral Ecology, 28,
642–650. https://doi.org/10.1046/j.1442-9993.2003.01321.x
Pompanon, F., Deagle, B. E., Symondson, W. O. C., Brown, D., Jarman,
S. N., & Taberlet, P. (2012). Who is eating what: Diet assessment
using next generation sequencing. Molecular Ecology, 21, 1931–1950.
https://doi.org/10.1111/j.1365-294X.2011.05403.x
Price, P. W. (2002). Resource‐driven terrestrial interac‐
tion webs. Ecological Research, 17, 241–247. https://doi.
org/10.1046/j.1440-1703.2002.00483.x
R Development Core and Team (2016). R: A language and environment
for statistical computing. Vienna, Austria: R Foundation for Statistical
Computing.
Ratajczak, Z., Nippert, J., & Ocheltree, T. W. (2014). Abrupt transition
of mesic grassland to shrubland: Evidence for thresholds, alternative
attractors, and regime shifts. Ecology, 95, 2633–2645. https://doi.
org/10.1890/13-1369.1
Rezende, E., Albert, E., Fortuna, M., & Bascompte, J. (2009).
Compartments in a marine food web associated with phylogeny,
body mass, and habitat structure. Ecology Letters, 12, 779–788.
https://doi.org/10.1111/j.1461-0248.2009.01327.x
Rowell‐Rahier, M. (1984). The presence or absence of phenolglycosides
in Salix (Salicaceae) leaves and the level of dietary specialisation of
some of their herbivorous insects. Oecologia, 62, 26–30. https://doi.
org/10.1007/BF00377368
Schleuning, M., Fründ, J., Schweiger, O., Welk, E., Albrecht, J., Albrecht,
M., … Hof, C. (2016). Ecological networks are more sensitive to plant
than animal extinction under climate change. Nature Communications,
7, 13965.
Schloss, P. D., Westcott, S. L., Ryabin, T., Hall, J. R., Hartmann, M.,
Hollister, E. B., ... Weber, C. F. (2009). Introducing mothur: Open-
source, platform-independent, community-supported software
for describing and comparing microbial communities. Applied and
Environmental Microbiology, 75, 7537–7541. https://doi.org/10.1128/
AEM.01541-09
Singer, M., & Stireman, J. (2001). How foraging tactics determine host‐
plant use by a polyphagous caterpillar. Oecologia, 129, 98–105.
https://doi.org/10.1007/s004420100707
Soininen, E. M., Valentini, A., Coissac, E., Miquel, C., Gielly, L., Brochmann,
C., … Taberlet, P. (2009). Analysing diet of small herbivores: The ef‐
ficiency of DNA barcoding coupled with high‐throughput pyrose‐
quencing for deciphering the composition of complex plant mixtures.
Frontiers in Zoology, 6, 16. https://doi.org/10.1186/1742-9994-6-16
Taberlet, P., Gielly, L., Pautou, G., & Bouvet, J. (1991). Universal primers
for amplification of three non‐coding regions of chloroplast DNA.
Plant Molecular Biology, 17, 1105–1109. https://doi.org/10.1007/
BF00037152
Thébault, E., & Fontaine, C. (2010). Stability of ecological communities
and the architecture of mutualistic and trophic networks. Science,
329, 853–856. https://doi.org/10.1126/science.1188321
Towne, E. G. (2002). Vascular plants of Konza Prairie Biological Station:
An annotated checklist of species in a Kansas tallgrass prairie. Sida,
20, 269–294.
Towne, E. G., Hartnett, D. C., & Cochran, R. C. (2005). Vegetation
trends in tallgrass prairie from bison and cattle grazing. Ecological
Applications, 15, 1550–1559. https://doi.org/10.1890/04-1958
WELTI et al. Functional Ecology |
      11

Tylianakis, J. M., Tscharntke, T., & Lewis, O. T. (2007). Habitat modifi‐ and their consequences for grassland plant diversity. Ecology, 95,
cation alters the structure of tropical host-parasitoid food webs. 1055–1064. https://doi.org/10.1890/13-1079.1
Nature, 455, 202–205. Zuur, A. F., Ieno, E. N., Walker, N. J., Saveliev, A. A., & Graham, M. S.
Ueckert, D. N., Hansen, R. M., & Terwilliger, C. Jr (1972). Influence of (2009). Mixed effects models and extensions in ecology with R. New
plant frequency and certain morphological variation on diets of York, NY: Springer‐Verlag.
rangeland grasshoppers. Journal of Range Management, 25, 61–65.
Unsicker, S. B., Specht, F. A., Köhler, G., Linz, J., Renker, C., Stein, C., &
Weisser, W. W. (2010). Plant species richness in montane grasslands
affects the fitness of a generalist grasshopper species. Ecology, 91, S U P P O R T I N G I N FO R M AT I O N
1083–1091. https://doi.org/10.1890/09-0402.1
Welti, E. A. R., Helzer, C., & Joern, A. (2017). Impacts of plant diversity on Additional supporting information may be found online in the
arthropod communities and plant‐herbivore network architecture. Supporting Information section at the end of the article.
Ecosphere, 8, e01983. https://doi.org/10.1002/ecs2.1983
Welti, E. A. R., & Joern, A. (2015). Structure of trophic and mutualis‐
tic networks across broad environmental gradients. Ecology and
How to cite this article: Welti EAR, Qiu F, Tetreault HM,
Evolution, 5, 326–334. https://doi.org/10.1002/ece3.1371
Welti, E. A. R., Qiu, F., Tetreault, H. M., Ungerer, M., Blair, J., & Joern, A. Ungerer M, Blair J, Joern A. Fire, grazing and climate shape
(2018). Data from: Fire, grazing, and climate shape plant‐grasshopper plant–grasshopper interactions in a tallgrass prairie. Funct
interactions in a tallgrass prairie. Dryad Digital Repository, https://doi. Ecol. 2019;00:1–11. https://doi.org/10.1111/1365-
org/10.5061/dryad.s6j1822
2435.13272
Zhong, Z., Wang, D., Zhu, H., Wang, L., Feng, C., & Wang, Z. (2014).
Positive interactions between large herbivores and grasshoppers,