Grasshopper Performance Research Articles

Woody vegetation remnants within pastures influence locust distribution: Testing bottom-up and top-down control

Agriculture is a major factor in landscape fragmentation, altering nutrient cycling and animal and plant populations through increasing habitat edge density. Most research on insect herbivores in agroecosystems has focused on the top-down effects of predators moving throughout the habitat mosaic. Few studies have focused on the top-down and bottom-up factors modulating the distribution of insect herbivore populations between natural and agricultural patches. For example, despite an understanding that Australian plague locusts (Chortoicetes terminifera) avoid tree patches, the underlying mechanisms remain unknown. Here, we explored how wooded remnants within pastures affect locust density and the potential top-down and bottom-up mechanistic explanations. We tested three hypotheses: 1) grasses near wooded areas are nutritionally suboptimal, 2) predator density is higher near wooded areas, and 3) temperatures are cooler underneath trees. We measured locust density, grass nutrient content, predator abundance, temperature, and ground cover along 50 m transects from wooded areas to open grassy areas. We ran those transects in three fields and had four transects per field. We confirmed locust avoidance for trees at a 20 m periphery, however none of the variables tested independently explained this trend. Grass nutrient content was similar underneath wooded areas and in open patches. Predator abundance did not differ between the two habitats. The ground was warmer under wooded areas than in grassy areas potentially due to woody vegetation negating windchill. Further, we found that locust density was negatively correlated with plant protein content and was highest in areas with approximately 20 % bare ground cover. Both plant protein and ground cover are important for grasshopper performance and reproduction. It is likely a complex interaction between these variables and others that drive the distribution of this species and other insect herbivores in agroecosystems. The small-scale mechanisms driving the response of insect herbivores to landscape changes is critical to understanding and predicting population dynamics at large-scales.

Plant composition changes in a small-scale community have a large effect on the performance of an economically important grassland pest

BackgroundThe grasshopper Oedaleus asiaticus Bey-Bienko (Acrididae: Oedipodinae) is a dominant and economically important pest that is widely distributed across the Mongolian plateau. This herbivore pest causes major damage to the grassland of the Inner Mongolian steppe in China. The population dynamics of herbivore pests is affected by grassland management practices (e.g., mowing and heavy livestock grazing) that alter plant community structures and stoichiometric characteristics. For example, O. asiaticus outbreak is closely associated with plant preference changes caused by nitrogen loss from heavy livestock grazing. However, the manner by which small-scale variation in vegetation affects grasshopper performance and promotes outbreak is poorly characterized. To address this question, we investigated the relationship between small-scale (1 m2) vegetation variability and measures of O. asiaticus performance associated with plant stoichiometric characteristics.ResultsWe found that food preferences of O. asiaticus varied significantly, but maintained a specific dietary structure for different plant compositions. Notably, small-scale changes in plant community composition significantly affected grasshopper food preference and body size. Partial least-square modeling indicated that plant proportion and biomass affected grasshopper body size and density. We found that this effect differed between sexes. Specifically, female body mass positively correlated with the proportion of Stipa krylovii grass, whereas male mass positively correlated with the proportion of Artemisia frigida grass. Further analyses indicated that grasshopper performance is closely associated with plant stoichiometric traits that might be responsible for the pest’s plague.ConclusionsThis study provides valuable information for managing grasshoppers using rational grassland management practices.

Molecular Ecological Basis of Grasshopper (Oedaleus asiaticus) Phenotypic Plasticity under Environmental Selection.

While ecological adaptation in insects can be reflected by plasticity of phenotype, determining the causes and molecular mechanisms for phenotypic plasticity (PP) remains a crucial and still difficult question in ecology, especially where control of insect pests is involved. Oedaleus asiaticus is one of the most dominant pests in the Inner Mongolia steppe and represents an excellent system to study phenotypic plasticity. To better understand ecological factors affecting grasshopper phenotypic plasticity and its molecular control, we conducted a full transcriptional screening of O. asiaticus grasshoppers reared in four different grassland patches in Inner Mongolia. Grasshoppers showed different degrees of PP associated with unique gene expressions and different habitat plant community compositions. Grasshopper performance variables were susceptible to habitat environment conditions and closely associated with plant architectures. Intriguingly, eco-transcriptome analysis revealed five potential candidate genes playing important roles in grasshopper performance, with gene expression closely relating to PP and plant community factors. By linking the grasshopper performances to gene profiles and ecological factors using canonical regression, we first demonstrated the eco-transcriptomic architecture (ETA) of grasshopper phenotypic traits (ETAGPTs). ETAGPTs revealed plant food type, plant density, coverage, and height were the main ecological factors influencing PP, while insect cuticle protein (ICP), negative elongation factor A (NELFA), and lactase-phlorizin hydrolase (LCT) were the key genes associated with PP. Our study gives a clear picture of gene-environment interaction in the formation and maintenance of PP and enriches our understanding of the transcriptional events underlying molecular control of rapid phenotypic plasticity associated with environmental variability. The findings of this study may also provide new targets for pest control and highlight the significance of ecological management practice on grassland conservation.

Effects of Altered Seasonality of Precipitation on Grass Production and Grasshopper Performance in a Northern Mixed Prairie.

Climatic changes are leading to differing patterns and timing of precipitation in grassland ecosystems, with the seasonal timing of precipitation affecting plant biomass and plant composition. No previous studies have examined how drought seasonality affects grasshopper performance and the impact of herbivory on vegetation. We modified seasonal patterns of precipitation and grasshopper density in a manipulative experiment to examine if seasonality of drought combined with herbivory affected plant biomass, nitrogen content, and grasshopper performance. Grass biomass was affected by both precipitation and grasshopper density treatments, while nitrogen content of grass was higher with early-season drought. Proportional survival was negatively affected by initial density, while survival was higher with early drought than with full-season drought. Drought timing affected the outcome, with early summer drought increasing grass nitrogen content and grasshopper survival, while season-long and late-season drought did not. The results support arguments that our knowledge of plant responses to seasonal short-term variation in climate is limited and illustrate the importance of experiments manipulating precipitation phenology. The results confirm that understanding the season of drought is critical for predicting grasshopper population dynamics, as extreme early summer drought may be required to strongly affect Melanoplus sanguinipes (F.) performance.

Abiotic effects on the clear-winged grasshopper (Orthoptera: Acridae) and its fungal pathogen Entomophaga grylli (Entomophthorales: Entomophthoraceae) in an intermountain bunch-grass prairie

AbstractThe grasshopper, Camnula pellucida Scudder (Orthoptera: Acridae), is a severe pest of small grains and rangeland forage in North America. In a field experiment using cages containing C. pellucida in northwestern Montana, United States of America, we manipulated exposure to the fungal entomopathogen, Entomophaga grylli Fresenius (Entomophthorales: Entomophthoraceae) pathotype 1, temperature using small greenhouses, and moisture for the pathogen with water pillows. Treatment effects on fungal infection (mycosis) rates and grasshopper survivorship were assessed. Water pillows provided additional moist habitat for E. grylli without having an impact on grasshopper performance or the plant biomass and nitrogen content. Number of fungal spores, pathogen-induced mortality rates, and treatment effects on grass biomass and nitrogen content were also measured. Water pillows benefited the pathogen by increasing fungal spore levels by eightfold, which in turn shortened pathogen-exposed grasshopper survival time by half. In contrast, warming reduced fungal spore levels by 50% and subsequently reduced mortality from E. grylli by 67%. However, warmed pathogen-exposed grasshoppers did not exhibit enhanced survival, which may be due, in part, to intraspecific competition among the surviving grasshoppers.

Density mediates grasshopper performance in response to temperature manipulation and spider predation in tallgrass prairie.

Species interactions are often context-dependent, where outcomes require an understanding of influences among multiple biotic and abiotic factors. However, it remains unclear how abiotic factors such as temperature combine with important biotic factors such as density-dependent food limitation and predation to influence species interactions. Using a native grassland - grasshopper - wolf spider model food chain in tallgrass prairie, we conducted a manipulative field experiment to examine how predator-prey interactions respond to manipulations of temperature, grasshopper density, and food chain length. We find that grasshopper performance responses to temperature and predator treatments were density dependent. At high densities, grasshopper survival decreased with increased temperature when no spiders were present. When spiders were present, grasshopper survival was reduced, and this effect was strongest in the cooled treatment. In contrast, grasshopper survival did not vary significantly with spider presence or among temperature treatments at low grasshopper densities. Our results indicate that context-dependent species interactions are common and highlight the importance of understanding how and when key biotic and abiotic factors combine to influence species interactions.

Herbivory of native and exotic North-American prairie grasses by nymph Melanoplus grasshoppers

Although the general interaction between native and exotic plants and specialist insect herbivores has received considerable attention in ecological studies, plant responses to herbivory by generalist insects, as well as feeding preferences of generalist insects, are still poorly understood. Experimental studies comparing leaf damage of native and exotic plants to generalist insects have provided inconsistent results due to irregularities in methodology such as using only one type of experiment or using non-standard measurements. Our study addresses these issues by examining leaf damage caused by generalist Melanoplus grasshoppers in morphologically and physiologically similar native (Andropogon gerardii and Bouteloua curtipendula) and exotic grasses (Miscanthus sinensis and Bothriochloa ischaemum) in Ohio and Maryland. In a 2-year study, we explored whether exotic grasses sustain less damage than native grasses, and consequently, whether generalist grasshoppers may potentially pose biotic resistance to exotic grasses. Using a combination of choice and no-choice feeding experiments with intact plants and with clipped leaves under different (field and greenhouse) conditions, we found that overall both herbivory and grasshopper performance (body length) were greater on the exotic grasses compared to native grasses. Our results suggest that exotic grasses which do not share a coevolutionary history with native generalist Melanoplus grasshoppers might have lower physical and chemical defenses than native plants. Consequently, Melanoplus grasshoppers may provide biotic resistance to these exotic grasses should these plants invade natural areas at the study regions. These results have important applications for predicting the interaction between exotic plants and generalist herbivores in the introduced range, which is critical for understanding factors facilitating plant invasions.

Performance and Feeding Preference ofCornops aquaticum(Orthoptera: Acrididae) onEichhornia crassipesand Crop Plants in Native Area

Abstract The semiaquatic grasshopper Cornops aquaticum Bruner, which is highly specific of Eichhornia crassipes (Pontederiaceae), constitutes one of the most abundant herbivores in the Neotropical wetlands and a possible biological control agent of this weed in South Africa. The aims of this study were 1) to assess the performance of each age category of C. aquaticum on crop plants and the host plant E. crassipes; 2) to assess the feeding preference of this grasshopper on crop plants in the presence and absence of E. crassipes; and 3) to determine the presence of nymphs and adults of this grasshopper on crops and in the adjacent wetlands where Eichhornia plants are present. Crop plants included in this study were Zea mays, Oryza sativa, Saccharum officinarum and Ilex paraguariensis. The age categories of C. aquaticum population were nymphs A (instar I and II), nymphs B (instar III to VI) and adults (females and males). To assess grasshopper performance on E. crassipes and crop plants, survival and oviposi...

Increase of fast nutrient cycling in grassland microcosms through insect herbivory depends on plant functional composition and species diversity

Nutrient cycling in terrestrial ecosystems is affected by various factors such as plant diversity and insect herbivory. While several studies suggest insect herbivory to depend on plant diversity, their interacting effect on nutrient cycling is unclear.In a greenhouse experiment with grassland microcosms of one to six plant species of two functional groups (grasses and legumes), we tested the influence of plant species richness (diversity) and functional composition on plant community biomass production, insect foliar herbivory, soil microbial biomass, and nutrient concentrations in throughfall. To manipulate herbivory, zero, three or six generalist grasshoppers (Chorthippus parallelus) were added to the plant communities.Increasing plant species richness increased shoot biomass and grasshopper performance, without significantly affecting root biomass or insect herbivory. Plant functional composition affected all of these parameters, e.g. legume communities showed the highest shoot biomass, the lowest grasshopper performance and suffered the least herbivory. Nutrient concentrations (dissolved mineral N, PO4‐P, SO4‐S) and pH in throughfall increased with herbivory. PO4‐P and pH increases were positively affected by plant diversity, especially under high herbivore pressure. Plant functional composition affected several throughfall variables, sometimes fully explaining diversity effects. Increasing plant diversity tended to increase soil microbial biomass, but only under high herbivore pressure. Faeces quantities strongly correlated with changes in pH and PO4‐P; frass may therefore be an important driver of throughfall pH and a main source of PO4‐P released from living plants. Our results indicate that insect herbivory may significantly influence fast nutrient cycling processes in natural communities, particularly so in managed grasslands.

Grasshoppers amensalistically suppress caterpillar performance and enhance plant biomass in an alpine meadow

Amensalism may be common between non‐trophically linked animals in natural ecosystems, where variation among species in body sizes and foraging modes may give rise to one‐sided interference. However, species and ecosystem‐level consequences of animal–animal amensalism are largely unknown. In a Tibetan alpine meadow, dominant herbivorous grasshoppers trigger a death feigning anti‐predator response of co‐occurring grassland caterpillars despite posing no consumptive threat. We hypothesized that: 1) grasshoppers reduce the performance of caterpillars while incurring no cost to themselves; and 2) this amensalism reduces top–down control of plant composition and biomass. We tested these hypotheses by factorial manipulation of both herbivores within replicate field enclosures. Grasshoppers significantly suppressed caterpillar feeding, growth rate, survival, reproductive effort and delayed metamorphosis. In contrast, grasshopper performance was unaffected by the caterpillars. Suppression of caterpillar feeding decreased overall herbivore suppression of plant biomass by 58% and shifted the functional composition of the plant community (i.e. increased sedge: forb ratio). These results suggest that consideration of non‐trophic interactions such as amensalism will help predict the consequences of species losses for the structure and functioning of ecosystems.

Predator–prey interactions in a grassland food chain vary with temperature and food quality

Because species interactions are often context‐dependent, abiotic factors such as temperature and biotic factors such as food quality may alter species interactions with potential consequences to ecosystem structure and function. For example, altered predator–prey interactions may influence the dynamics of trophic cascades, affecting net primary production. In a three‐year field experiment, we manipulated a plant–grasshopper–spider food chain in mesic tallgrass prairie to investigate the effects of temperature and food quality on grasshopper performance, and to understand the direct and indirect tritrophic interactions that contribute to trophic cascades. Because spiders are active at cooler temperatures than grasshoppers in our system, we hypothesized that predator effects would be strongest in cooled treatments, and weakest in warmed treatments. Grasshopper spider interactions were highly context‐dependent and varied significantly with food quality, temperature treatment and year. Spiders most often reduced grasshopper survival in the cooled and ambient temperature treatments, but had little to no effect on grasshopper survival in the warmed treatments, as hypothesized. In some years, plants compensated for grasshopper herbivory and trophic cascades were not observed despite significant effects of predators on grasshopper survival. However, in the year they were observed, trophic cascades only occurred in cooled treatments where predator effects on grasshoppers were strongest. Predicting ecosystem responses to climate change will require an understanding of how temperature influences species interactions. Our results demonstrate that changes in daily temperature regimes can alter predator–prey interactions among arthropods with consequences for ecosystem processes such as primary production and the relative importance of top–down and bottom–up processes.

Stressing food plants by altering water availability affects grasshopper performance

Extreme weather events like drought and heavy rain are likely to increase with climate change in Central Europe and may affect nutrient content in plants. They may therefore influence the performance (growth rate, developmental time, mortality, body size/mass, fecundity) and population dynamics of herbivorous insects. We conducted a common-garden experiment on food plants to investigate effects of severe drought and moisture events on reproduction and fitness components in the insect herbivore Chorthippus biguttulus (Orthoptera, Acrididae). Periodic irrigations of food plants were used to simulate a 60% decrease in average summer precipitation (drought treatment), a 40% increase in average summer precipitation (moisture treatment) and a normal summer precipitation (control treatment). Individuals of C. biguttulus that fed on drought-stressed plants showed beneficial effects on life-history traits including an increased reproductive success than grasshoppers that fed on control plants. The opposite was tru...

How Will Species Respond to Climate Change? Examining the Effects of Temperature and Population Density on an Herbivorous Insect

An important challenge facing ecologists is to understand how climate change may affect species performance and species interactions. However, predicting how changes in abiotic variables associated with climate change may affect species performance also depends on the biotic context, which can mediate species responses to climatic change. We conducted a 3-yr field experiment to determine how the herbivorous grasshopper Camnula pellucida (Scudder) responds to manipulations of temperature and population density. Grasshopper survival and fecundity decreased with density, indicating the importance of intraspecific competition. Female fecundity tended to increase with temperature, whereas grasshopper survival exhibited a unimodal response to temperature, with highest survival at intermediate temperatures. Grasshopper performance responses to temperature also depended on density. Peak survival in the low-density treatment occurred in warmer conditions than for the high-density treatment, indicating that the intensity of intraspecific competition varies with temperature. Our data show that changes to the temperature regimen can alter grasshopper performance and determine the intensity of intraspecific competition. However, the effects of temperature on grasshopper performance varied with density. Our data indicate the importance of the biotic context in mediating species responses to climatic factors associated with global change.

Resource-mediated impact of spider predation risk on performance in the grasshopper Ageneotettix deorum (Orthoptera: Acrididae).

In response to increased exposure to predators when searching for food, many prey increase the frequency of antipredator behaviors, potentially reducing foraging rate and food intake. Such direct, nonlethal interactions between predators and prey resulting in reduced food intake can indirectly influence lifecycle development through effects on growth, developmental rate, and survival. We investigated the general hypothesis that individual performance of a herbivorous insect can be negatively affected when exposed to nonlethal predation risk, and that the response can be mediated by food quality. This hypothesis was tested using the common rangeland grasshopper Ageneotettix deorum with and without exposure to common wolf spider predators (Lycosidae, Schizocosa spp.) on both untreated natural and fertilized vegetation. All spiders were rendered temporarily incapable of direct feeding by restricting function of the chelicerae with beeswax. Detectable responses by grasshoppers to spiders indicate indirect consequences for lifecycle development. Grasshopper performance was measured as hind femur growth, duration of nymphal lifecycle stages, and survivorship in a caged field experiment conducted over 2 years. Grasshoppers developed faster and grew 3-5% larger when allowed to forage on fertilized vegetation in the absence of risk from a spider predator. Failure-time analysis illustrated enhanced survival probability in response to elevated food quality and the negative effects of grasshopper susceptibility to nonlethal predation risk. Performance on food of relatively low, ambient quality with no predation risk equaled that of grasshoppers caged with high quality vegetation in the presence of a modified spider. Increased resource quality can clearly moderate the negative life history responses caused by the behavioral modification of grasshoppers when exposed to spider predation risk, a compensatory response.