Herbivore phenology can predict response to changes in plant quality by livestock grazing

Livestock grazing can have a strong impact on herbivore abundance, distribution and community. However, not all species of herbivores respond the same way to livestock grazing, and we still have a poor understanding of the underlying mechanisms driving these differential responses. Here, we investigate the effect of light intensity cattle grazing on the abundance of two grasshoppers (Euchorthippus cheui and E. unicolor) that co-occur in the same grasslands and feed on the same food plant (the dominant grass Leymus chinensis). The two grasshopper species differ in phenology so that their peak abundances are separated into early- and late-growing seasons. We used an exclosure experiment to monitor grasshopper abundance and food quality in the field under grazed and ungrazed conditions, and performed feeding trials to examine grasshopper preference for grazed or ungrazed food plants in the laboratory. We found that the nitrogen content of L. chinensis leaves continuously declined in the ungrazed areas, but was significantly enhanced by cattle grazing over the growing season. Cattle grazing facilitated the early-season grasshopper E. cheui, whereas it suppressed the late-season grasshopper E. unicolor. Moreover, feeding trials showed that E. cheui preferred L. chinensis from grazed plots, while E. unicolor preferred the leaves from ungrazed plots. We conclude that livestock grazing has opposite effects on the two grasshopper species, and that these effects may be driven by grazing-induced changes in plant nutrient content and the unique nutritional niches of the grasshoppers. These results suggest that insects that belong to the same guild can have opposite nutrient requirements, related to their distinct phenologies, and that this can ultimately affect their response to cattle grazing. Our results show that phenology may link insect physiological needs to local resource availabilities, and should be given more attention in future work on interactions between large herbivores and insects.     

Impacts of grazing intensity and increased precipitation on a grasshopper assemblage (Orthoptera: Acrididae) in a meadow steppe

1. Grasshoppers are dominant herbivores in grassland ecosystems, and many studies have examined how grazing by large herbivores and precipitation patterns individually influence the dynamics of grassland grasshopper assemblages, but their combined effects are largely unknown. 2. In this study, grazing intensities (ungrazed, moderate, and heavy) were manipulated and precipitation (ambient and increased amount of rainfall) altered in a field experiment to test the effects of grazing and altered precipitation on a grasshopper community in a meadow steppe in northeastern China. 3. It was found that grasshopper species richness did not change according to different grazing intensities under ambient precipitation, but was significantly higher (by 38.1%) in moderate grazing intensities under increased precipitation. Grasshopper abundance increased considerably with increasing grazing intensities in ambient precipitation treatments; however, grasshopper abundance in heavy grazing intensities was significantly lower (by 32.9%) than in the other two grazing intensities under increased precipitation. Moreover, the responses of grasshopper abundance to grazing under altered precipitation were species‐specific. 4. Grazing effects on grasshopper species diversity were mediated through the species richness and biomass of grasses (food resources), but the effects on grasshopper abundance were mediated through plant height (vegetation structure) under altered precipitation. 5. These results suggest that appropriate grazing by large herbivores would be considered as beneficial management practices for maintaining grasshopper diversity and abundance under conditions of increased precipitation in grassland ecosystems. Additionally, greater attention should be paid to the population dynamics of different grasshopper species to better understand the responses of grasslands to grazing and altered precipitation.     

Large herbivores facilitate an insect herbivore by modifying plant community composition in a temperate grassland

Large herbivores often co‐occur and share plant resources with herbivorous insects in grassland ecosystems; yet, how they interact with each other remains poorly understood. We conducted a series of field experiments to investigate whether and how large domestic herbivores (sheep; Ovis aries) may affect the abundance of a common herbivorous insect (aphid; Hyalopterus pruni) in a temperate grassland of northeast China. Our exclosure experiment showed that 3 years (2010–2012) of sheep grazing had led to 86% higher aphid abundance compared with ungrazed sites. Mechanistically, this facilitative effect was driven by grazing altering the plant community, rather than by changes in food availability and predator abundance for aphids. Sheep significantly altered plant community by reducing the abundance of unpalatable forbs for the aphids. Our small‐scale forb removal experiment revealed an “associational plant defense” by forbs which protect the grass Phragmites australis from being attacked by the aphids. However, selective grazing on forbs by sheep indirectly disrupted such associational plant defense, making P. australis more susceptible to aphids, consequentially increasing the density of aphids. These findings provide a novel mechanistic explanation for the effects of large herbivores on herbivorous insects by linking selective grazing to plant community composition and the responses of insect populations in grassland ecosystems.     

Insect herbivory and vertebrate grazing impact food limitation and grasshopper populations during a severe outbreak

1. Competition between herbivores often plays an important role in population ecology and appears strongest when densities are high or plant production is low. Phytophagous insects are often highly abundant, but relatively few experiments have examined competition between vertebrates and phytophagous insects. 2. In grassland systems worldwide, grasshoppers are often the dominant phytophagous insect, and livestock grazing is a dominant land use. For this study, a novel experiment was conducted examining competition between vertebrates and invertebrates, where both grasshopper densities and sheep grazing were manipulated inside 10‐m² caged mesocosms during a grasshopper outbreak. We examined how grasshopper densities and the timing of vertebrate herbivory affected grasshopper densities, if the effects of vertebrates on survival and reproduction changed with grasshopper density, and how a naturally occurring grasshopper outbreak affected grasshopper populations in the following year. 3. Densities of grasshoppers at the site peaked at 130 m^–2. Food‐limited competition was stronger in treatments with higher grasshopper densities and repeated or late livestock herbivory, leading to reduced survival, femur length, and functional ovarioles, a measure of future reproduction. Strong food‐limited density‐dependent reproduction and survival led to reduced hatching densities in 2001. 4. As competition was typically stronger with high grasshopper densities than with livestock grazing, competition from vertebrates could be relatively less important for phytophagous insect population dynamics during outbreaks. The experiment provides insights into how competition between insect and vertebrate herbivores influences insect population dynamics, and indicates that severe outbreaks can rapidly subside with strong competition from vertebrate and insect herbivores.     

Grasshopper (Orthoptera: Acrididae) communities respond to fire,bison grazing and weather in North American tallgrass prairie: a long-term study

Because both intrinsic and extrinsic factors influence insect population dynamics, operating at a range of temporal and spatial scales, it is difficult to assess their contributions. Long-term studies are ideal for assessing the relative contributions of multiple factors to abundance and community dynamics. Using data spanning 25 years, we investigate the contributions of weather at annual and decadal scales, fire return interval, and grazing by bison to understand the dynamics of abundance and community composition in grasshopper assemblages from North American continental grassland. Each of these three primary drivers of grassland ecosystem dynamics affects grasshopper population and community dynamics. Negative feedbacks in abundances, as expected for regulated populations, were observed for all feeding guilds of grasshoppers. Abundance of grasshoppers did not vary in response to frequency of prescribed burns at the site. Among watersheds that varied with respect to controlled spring burns and grazing by bison, species composition of grasshopper assemblages responded significantly to both after 25 years. However, after more than 20 years of fire and grazing treatments, the number of years since the last fire was more important than the managed long-term fire frequency per se. Yearly shifts in species composition (1983–2005), examined using non-metric multidimensional scaling and fourth-corner analysis, were best explained by local weather events occurring early in grasshopper life cycles. Large-scale patterns were represented by the Palmer Drought Severity Index and the North Atlantic Oscillation (NAO). The NAO was significantly correlated with annual mean frequencies of grasshoppers, especially for forb- and mixed-feeding species. Primary grassland drivers—fire, grazing and weather—contributing both intrinsic and extrinsic influences modulate long-term fluctuations in grasshopper abundances and community taxonomic composition. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Cattle grazing in CRP grasslands during the nesting season: effects on avian abundance and diversity

The Conservation Reserve Program (CRP) is a primary tool for restoring grassland in the United States, in part as wildlife habitat, which has benefited declining grassland bird populations. Among potential mid-contract management practices used to maintain early-successional CRP grasslands, cattle grazing had been prohibited and is currently disincentivized during the primary nesting season for birds (much of the growing season), despite the important role that large herbivores historically played in structuring grassland ecosystems. Conservative grazing of CRP grasslands could increase spatial heterogeneity in vegetation structure and plant diversity, potentially supporting higher densities of some grassland bird species and higher bird diversity. Our objective was to determine the effect of experimental cattle grazing on species-specific relative abundance and occupancy, species diversity, and community dissimilarity of grassland birds on CRP grasslands across the longitudinal extent of Kansas, USA (a 63.5-cm precipitation gradient) during the 2017–2019 avian breeding seasons. Fifty-three of 108 fields were grazed by cattle during the growing seasons of 2017 and 2018 and all fields were rested from grazing in 2019. For all analyses, we examined separate model sets for semiarid western versus more mesic eastern Kansas. Using data from line transect surveys, we modeled relative abundances of 5 songbird species: grasshopper sparrow (Ammodramus savannarum), dickcissel (Spiza americana), eastern meadowlark (Sturnella magna), western meadowlark (Sturnella neglecta), and brown-headed cowbird (Molothrus ater). Grazing had delayed yet positive effects on abundances of grasshopper sparrow in western Kansas, and eastern meadowlark in eastern Kansas, but negative effects on dickcissel abundance in western Kansas and especially on burned fields in eastern Kansas. Somewhat counterintuitively, brown-headed cowbirds in western Kansas were more abundant on ungrazed versus grazed fields in the years after grazing began. In addition, we modeled multi-season occupancy of 3 gamebird species (ring-necked pheasant [Phasianus colcicus], northern bobwhite [Colinus virginianus], mourning dove [Zenaida macroura]) and Henslow's sparrow (Centronyx henslowii); grazing did not affect occupancy of these species. In eastern Kansas, species diversity was highest in grazed, unburned fields. In western Kansas, bird communities in grazed and ungrazed fields were dissimilar, as determined from multivariate analysis. Though regionally variable, conservative stocking of cattle on CRP grasslands during the nesting season as a mid-contract management tool might increase bird species diversity by restructuring habitat that accommodates a greater variety of species and decreasing abundances of species associated with taller, denser stands of vegetation.     

Being a generalist herbivore in a diverse world: how do diets from different grasslands influence food plant selection and fitness of the grasshopper Chorthippus parallelus?

1. Generalist insect herbivores occupy a variety of habitats that differ in food plant composition. Dietary mixing has been proposed as a possibility for generalists to overcome nutritional deficiencies of single plant species, but only a few studies have investigated herbivore feeding and fitness for diets that resemble natural scenarios. We studied feeding behaviour, survival, and reproduction of the generalist grasshopper Chorthippus parallelus raised on food plants of four typical habitats. 2. Grasshopper diet consisted of grasses (92.5%), legumes (6.7%) and, in small quantities, other forbs (0.8%). Diet selection differed between the four food plant mixtures, and depended on grasshopper sex and developmental stage. There was no correlation between the relative abundance of plant species in the field and the fraction of these species in the grasshopper diet. 3. Grasshoppers survived on average for 40.4 ± 1.0 days before maturity, grew 106.8 mg until maturity moult, and females laid 4.1 ± 0.4 egg pods, each of which contained 8.5 ± 0.4 eggs. However, despite the differences in feeding behaviour, grasshopper fitness was the same in all of the four food plant mixtures. While the digestibility of ingested food was similar in the four different treatments, indices indicated differences in the conversion efficiency to body mass. 4. Our results show that C. parallelus is a plastic feeder with no fixed preferences in diet composition. The results emphasise that generalist herbivores can counteract putative quality deficiencies of single food plants by selective dietary mixing.     

放牧对草地昆虫多样性的影响研究进展

在草地生态系统中,大型草食动物放牧是重要的管理方式之一,对草地生物多样性起着关键的驱动作用。昆虫是草地生态系统中生物多样性的重要组成成分,对生态系统的食物网结构以及其功能与稳定性起着关键作用。已有研究结果表明,大型草食动物与昆虫存在密切联系,放牧对草地昆虫多样性或有正向、或负向、或无明显作用,这依赖于放牧管理方式、昆虫类群以及草地类型。放牧必然通过直接(采食、践踏或粪尿)或间接(植物群落组成或植被结构)作用对昆虫多样性产生显著的影响。当前,关于大型草食动物放牧对草地昆虫多样性影响研究较多,但是,从研究系统性、深入性和延续性来说还存在一定问题。本文在综述国内外对放牧对草地昆虫多样性的影响研究基础上,提出了今后的研究方向,对于理解放牧管理的草地昆虫多样性变化规律,以及为积极探索维持草地昆虫多样性的长期有效的科学管理措施提供理论指导。     

Plant functional traits reveal the relative contribution of habitat and food preferences to the diet of grasshoppers

Food preferences and food availability are two major determinants of the diet of generalist herbivores and of their spatial distribution. How do these factors interact and eventually lead to diet differentiation in co-occurring herbivores? We quantified the diet of four grasshopper species co-occurring in subalpine grasslands using DNA barcoding of the plants contained in the faeces of individuals sampled in the field. The food preferences of each grasshopper species were assessed by a choice (cafeteria) experiment from among 24 plant species common in five grassland plots, in which the four grasshoppers were collected, while the habitat was described by the relative abundance of plant species in the grassland plots. Plant species were characterised by their leaf economics spectrum (LES), quantifying their nutrient vs. structural tissue content. The grasshoppers’ diet, described by the mean LES of the plants eaten, could be explained by their plant preferences but not by the available plants in their habitat. The diet differed significantly across four grasshopper species pairs out of six, which validates food preferences assessed in standardised conditions as indicators for diet partitioning in nature. In contrast, variation of the functional diversity (FD) for LES in the diet was mostly correlated to the FD of the available plants in the habitat, suggesting that diet mixing depends on the environment and is not an intrinsic property of the grasshopper species. This study sheds light on the mechanisms determining the feeding niche of herbivores, showing that food preferences influence niche position whereas habitat diversity affects niche breadth.     

Tracking nutrients in space and time: Interactions between grazing lawns and drought drive abundances of tallgrass prairie grasshoppers

1. We contrast the response of arthropod abundance and composition to bison grazing lawns during a drought and non‐drought year, with an emphasis on acridid grasshoppers, an important grassland herbivore.
2. Grazing lawns are grassland areas where regular grazing by mammalian herbivores creates patches of short‐statured, high nutrient vegetation. Grazing lawns are predictable microsites that modify microclimate, plant structure, community composition, and nutrient availability, with likely repercussions for arthropod communities.
3. One year of our study occurred during an extreme drought. Drought mimics some of the effects of mammalian grazers: decreasing above‐ground plant biomass while increasing plant foliar percentage nitrogen.
4. We sampled arthropods and nutrient availability on and nearby (“off”) 10 bison‐grazed grazing lawns in a tallgrass prairie in NE Kansas. Total grasshopper abundance was higher on grazing lawns and the magnitude of this difference increased in the wetter year of 2019 compared to 2018, when drought led to high grass foliar nitrogen concentrations on and off grazing lawns. Mixed‐feeding grasshopper abundances were consistently higher on grazing lawns while grass‐feeder and forb‐feeder abundances were higher on lawns only in 2019, the wetter year. In contrast, the abundance of other arthropods (e.g., Hemiptera, Hymenoptera, and Araneae) did not differ on and off lawns, but increased overall in 2019, relative to the drought of 2018.
5. Understanding these local scale patterns of abundances and community composition improves predictability of arthropod responses to ongoing habitat change.

     

The effect of horse, cattle and sheep grazing on the diversity and abundance of land snails in nutrient-poor calcareous grasslands

Livestock grazing is a common management practise in semi-natural grasslands in Central Europe. Different types of livestock (horses, cattle, sheep) and grazing intensity are known to affect the richness and composition of plant species. However, knowledge of grazing-dependent effects on invertebrates is limited. We examined the influence of horse, cattle and sheep grazing on the richness, abundance and composition of land snail species in 21 calcareous nutrient-poor grassland areas in the northwestern Jura Mountains, Switzerland. Grazing by different livestock species did not affect the species richness, abundance and species composition of land snails. Furthermore, the number of open-land species and the ratio of large- to small-sized snail species or individuals did not differ among the three pasture types. However, independent of livestock species, grazing intensity negatively influenced the snail fauna. Snail species richness, abundance and number of Red list species decreased with increasing grazing intensity. Grazing intensity also affected the occurrence of individual snail species (Truncatellina cylindrica, Cecilioides acicula, Candidula unifasciata and Trichia plebeia). To preserve the snail fauna in nutrient-poor grasslands, pastures can be stocked with horses, cattle or sheep. However, both maximum stocking rate (number of livestock units per hectare) and grazing duration (number of grazing days per year) must be carefully defined for the proper management of the pastures.     

Patch dynamics in grazed subtropical native pastures in south‐east Queensland

Abstract Patch formation is common in grazed grasslands but the mechanisms involved in the formation and maintenance of patches are not clear. To increase our knowledge on this subject we examined possible reasons for patch formation and the influence of management on changes between patch states in three experiments in native pasture communities in the Crows Nest district, south‐east Queensland. In these communities, small‐scale patches (tall grassland (dominated by large and medium tussock grasses), short swards (dominated by short tussock grasses and sedges), and lawns (dominated by stoloniferous and/or rhizomatous grasses)) are readily apparent. We hypothesized that the formation of short sward and lawn patches in areas of tall grassland was due to combinations of grazing and soil fertility effects. This was tested in Experiment 1 by applying a factorial combination of defoliation, nutrient application and transplants of short tussock and stoloniferous species to a uniform area of tall grassland. Total species density declined during the experiment, was lower with high nutrient applications, but was not affected by defoliation. There were significant changes in abundance of species that provided support for our hypotheses. With light defoliation and low nutrients, the tall grassland remained dominated by large tussock grasses and contained considerable amounts of forbs. With heavy defoliation, the pastures were dominated by medium tussock grasses and there were significant decreases in forbs and increases in sedges (mainly with low nutrients) and stoloniferous grasses (mainly with high nutrients). Total germinable seed densities and those of most species groups were significantly lower in the heavy defoliation than the light defoliation plots. Total soil seed numbers were not affected by nutrient application but there were fewer seeds of the erect forbs and more sedge seeds in plots with high nutrients. The use of resting from grazing and fire to manage transitions between patches was tested. In Experiment 2 , changes in species density and abundance were measured for 5 years in the three patch types with and without grazing. Experiment 3 examined the effects of fire, grazing and resting on short sward patches over 4 years. In Experiment 2 , total species density was lower in lawn than short sward or tall grassland patches, and there were more species of erect forbs than other plant groups in all patch types. The lawn patches were originally dominated by Cynodon spp. This dominance continued with grazing but in ungrazed patches the abundance of Cynodon spp. declined and that of forbs increased. In the short sward patches, dominance of short tussock grasses continued with grazing but in ungrazed plots their abundance declined while that of large tussock grasses increased. The tall grassland patches remained dominated by large and medium tussock species. In Experiment 3 , fire had no effect on species abundance. On the grazed plots the short tussock grasses remained dominant but where the plots were rested from grazing the small tussock grasses declined and the large tussock grasses increased in abundance. The slow and relatively small changes in these experiments over 4 or 5 years showed how stable the composition of these pastures is, and that rapid changes between patch types are unlikely.     

Grasshopper assemblage response to a restored national park (Mountain Zebra National Park, South Africa)

Twelve grassland sites were sampled inside and outside the Mountain Zebra National Park (MZNP), South Africa to assess changes in grasshopper assemblages to grazing by indigenous mammals inside the park compared to grazing by domestic cattle outside. The MZNP has been restored from cattle-grazed farmland to indigenous mammal parkland for 62 years. The number of grasshopper species and families inside the park was not significantly different from outside the park, but the number of individuals inside the park was significantly higher. Multivariate statistics did not reveal any strong site groupings based on simple inside/outside comparisons, but there were clear groupings of sites based on vegetation characteristics and other environmental variables. The park boundary, therefore, does not significantly determine grasshopper assemblages, although intensity of grazing does. The indigenous mammals inside the park had the same effect on grasshoppers as the domestic cattle outside, and it was the level of defoliation and trampling that was important rather than the type of mammal. Intensive livestock grazing and trampling leads to bush encroachment and reduction in grass cover and/or disappearance of several grass species. In response to this pressure, grasshopper populations dropped, with localized extirpation of some species. Vegetation composition and structure, particularly grass height and percentage cover, had a significant effect on grasshopper assemblages. The MZNP is thus an area of localized, elevated grasshopper abundance in comparison with the surrounding farms, and presumably represents a situation prior to the current, intensive farming activities. Such elevated grasshopper abundances are important for maintaining soil quality and hence ecological integrity of this landscape which is poor in organics and nitrogen. The MZNP could be viewed as a centre where species with high mobility may seek refuge from anthropogenic pressures. The MZNP also serves as a reference illustrating the differences between restored-through-natural-succession and anthropogenically disturbed habitats, and compares desirable with undesirable ecosystem changes for herbivorous invertebrates such as grasshoppers.     

Low intensity, mixed livestock grazing improves the breeding abundance of a common insectivorous passerine

Livestock grazing is a major driver of ecosystem change and has been associated with significant declines in various bird species in Britain and worldwide. However, there is little experimental evidence to show how grazing affects bird populations. We manipulated livestock densities in a replicated field experiment and found that mixed sheep and cattle grazing, at low intensity, improved the breeding abundance of a common upland passerine, the meadow pipit Anthus pratensis, after two years. Plots stocked with sheep alone (at high or low density) or not stocked at all held fewer pipit territories. Despite a year-on-year decline in pairs of meadow pipits in intensively grazed plots, we found no effect of sheep number on breeding abundance. Our results support the hypothesis that mixed species of herbivores generate greater heterogeneity in vegetation structure, which modifies prey availability, resulting in a greater abundance of birds. The results of our study should inform the management of grassland areas and enhance the abundance of some bird species, particularly in areas that have seen significant shifts from mixed livestock grazing to grazing dominated by single species of animals.     

高寒草甸物种多样性对藏系牧羊放牧行为的影响

为探究草原植物物种多样性对家畜放牧行为的影响及其机制,在青藏高原高寒草甸开展藏系牧羊轮牧试验,调查植被物种多样性,观察藏系牧羊采食速率、觅食速率和采食时间,并计算藏系牧羊日采食量。结果表明：两年间,植物物种丰富度与藏系牧羊采食速率呈显著正相关关系（P<0.05）;觅食速率、采食时间和日采食量对放牧率响应敏感（P<0.05）,呈夏秋增冬春减的趋势。植物Shannon-Wiener指数与藏系牧羊的采食时间显著负相关（P<0.05）;在暖季或8羊/hm²放牧率下植物Shannon-Wiener指数与藏系牧羊日采食量呈显著正相关（P<0.05）。Pilelou均匀度指数与藏系牧羊采食速率和采食时间显著负相关（P<0.05）;在暖季或8羊/hm²放牧率下Pilelou均匀度指数与藏系牧羊日采食量呈显著负相关（P<0.05）。植物物种丰富度对藏系牧羊放牧行为贡献较大,且放牧藏系牧羊的采食速率和采食时间比觅食速率和日采食量对植物物种丰富度响应更敏感,以用植物物种丰富度为自变量可以更好预测藏系牧羊放牧行为。放牧管理通过影响植被物种多样性从而进一步影响了藏系牧羊放牧行为。放牧行为不仅是评价草地营养价值和家畜生产力的关键指标,也是草地健康管理的基础。因此,明确草原植物物种多样性-藏系牧羊放牧行为的互作机制有助于更好的提高藏系牧羊地生产力,维护草原生态健康。     

Water stress in grasslands: dynamic responses of plants and insect herbivores

Global climate change is altering precipitation patterns. The effect of water stress on plant–herbivore interactions is poorly understood even though this is a primary ecological interaction that will be altered by climate change. This is especially true for grasslands where water is often limiting. In this study we manipulated water inputs in open grassland plots (1 m²) during a severe drought and assessed plant and insect herbivore responses. There were two watering treatments: ambient and supplemented. Supplemented plots received water weekly in amounts that mimicked average seasonal rainfall. For plants, we were interested in how water input affected protein and digestible carbohydrate content; previous studies predicted water stress would increase the concentration of these two nutrients. Grasshoppers are the dominant insect herbivores in grasslands and we assessed their responses to water inputs by measuring abundance and diversity. Previous studies suggested grasshoppers would prefer water‐stressed plots. Protein and carbohydrate content in bulk grass and forb samples, plus plant biomass and diversity, were measured monthly (May–August). Immediately prior to harvesting plant tissue, we counted and identified individual grasshoppers in each plot. Grass biomass was reduced with water stress, but macronutrient content and species diversity were unaffected. After three months water‐stressed forbs were less protein biased, and diverse, relative to watered forbs; forb biomass was indistinguishable between treatments. Grasshopper abundance and diversity were lower in water‐stressed plots as the season progressed. However, grasshopper‐feeding biology mattered: densities of mixed‐feeders and grass‐feeders, but not forb‐specialists, decreased over time in water‐stressed plots, but not in water supplemented plots. Our results demonstrate the importance of focusing on plant and insect herbivore functional groups and provide valuable new data that can be incorporated into models to explore the effects of global climate change in greater detail.     

Yaks and sheep trigger different changes in the grasshopper assemblages of the Qilian Mountains via differentially altering plant assemblages

1. The Qilian Mountains represent one of the key livestock‐raising grasslands in China. The two main herbivore species raised in this area – yaks and sheep – are of critical economical value. Grasshoppers compete with these animals for available nutrients, creating multifaceted relationships between livestock, grasshoppers and plants. A clear understanding of such relationships is lacking and is urgently needed to guide conservation efforts. 2. This study aims to document the effects of yak and sheep grazing on grasshopper assemblages and to elucidate the underlying mechanisms of such effects. 3. It is shown here that yaks and sheep impact grasshopper assemblages differently. Grasshopper assemblages exhibited lower density, biodiversity, richness, and evenness of distribution in yak‐grazed pastures than in grazing‐free grasslands. Sheep‐grazed pastures exhibited a dramatically divergent picture, with elevated density, biodiversity and richness, and a slightly decreased evenness of distribution. Grasshoppers were generally larger in grazed pastures than in grazing‐free grasslands, especially in yak‐grazed plots. 4. The present study suggests that differences between yak and sheep pastures in plant assemblage structure and plant traits are probably the underlying forces driving the differences in grasshopper assemblage structure and grasshopper traits, respectively. 5. The study shows that the grasshopper habitat indicator species differ between yak and sheep pastures, raising the possibility that such indicators can be used to monitor grassland usage and degradation in the Qilian Mountains. 6. These results provide novel insights into the dynamic interactions of common domesticated herbivore species, grasshoppers and plants in Qilian Mountains, which augment current knowledge and may ultimately lead to better conservation practices.     

Predictions of species interactions from consumer-resource theory: experimental tests with grasshoppers and plants

We tested the ability of consumer-resource theory to predict direct and indirect interactions among species, using an experimental system of insect herbivores and herbaceous plants. Specifically, we examined interactions among three species of grasshoppers (Melanoplus femur-rubrum, Spharagemon collare, andPhoetaliotes nebrascensis; Orthoptera, Acrididae) and herbaceous plants in experimental field cages placed over existing fertilized or unfertilized vegetation in a Minnesota old field. For the conditions inside these cages, we addressed whether (1) grasshopper diet predicted the presence of competition among grasshopper species, and (2) direct effects of grasshoppers on plants produced indirect interactions among plants, grasshoppers and soil nitrogen. Overall,M. femur-rubrum ate a greater proportion of forbs in cages, while the other two species ate primarily grasses. As expected, a pair of grasshopper species competed if they had similar diets. However, there were important exceptions that could be explained from observed indirect effects, although alternative explanations were also possible. First, all three grasshopper species significantly shifted their diets in the presence of other species, and these shifts occurred most often when competition was expected or occurred. Second, the two grassfeeding species reduced the biomass of the dominant grass (Schizachyrium scoparium) and increased available soil nitrogen and biomass of forbs. This effect may explain why the grass-feedingP. nebrascensis had a positive effect on the forb-feedingM. femur-rubrum on unfertilized plots. Overall, we show that direct effects of consumers on resources can predict competition and other important indirect interactions within a community.     

Grasshopper fecundity responses to grazing and fire in a tallgrass prairie

Grasshopper abundance and diversity vary with management practices such as fire and grazing. Understanding how grasshopper life history traits such as fecundity respond to management practices is key to predicting grasshopper population dynamics in heterogeneous environments. Landscape-level experimental fire and bison grazing treatments at the Konza Prairie Biological Station (Manhattan, KS) provide an opportunity to examine how management affects grasshopper fecundity. Here we report on grasshopper fecundity for nine common species at Konza Prairie. From 2007 to 2009, adult female grasshoppers were collected every 3 wk from eight watersheds that varied in fire and grazing treatments. Fecundity was measured by examining female reproductive tracts, which contain a record of past and current reproductive activity. Body size was a poor predictor of fecundity for all species. Despite large differences in vegetation structure and composition with management regime (grazing and fire interval), we observed little effect of management on grasshopper fecundity. Habitat characteristics (grasshopper density, vegetation biomass, and vegetation quality; measured in 2008 and 2009) were better predictors of past fecundity than current fecundity, with species-specific responses. Fecundity increased throughout the summer, indicating that grasshoppers were able to acquire sufficient nutritional resources for egg production in the early fall when vegetation quality is generally low. Because fecundity did not vary across management treatments, population stage structure may be more important for determining population level reproduction than management regime at Konza Prairie.     

Foraging behaviour at multiple temporal scales in a wild alpine equid

Forage abundance, forage quality, and social factors are key elements of the foraging ecology of wild herbivores. For non-ruminant equids, forage-limited environments are likely to impose severe constraints on their foraging behaviour. We used a multi-scale approach to study foraging behaviour in kiang (Equus kiang), a wild equid inhabiting the high-altitude rangelands of the Tibetan Plateau. Using behavioural observations and vegetation sampling, we first assessed how patterns of plant abundance and quality affected (i) the instantaneous forage intake rate (fine scale) and (ii) the proportion of time spent foraging (coarse scale) across seasons. We also tested whether foraging behaviour differed among group types, between sex in adults, and between females of different reproductive status. At a fine scale, intake rate increased linearly with bite size and increased following a type II curvilinear function with biomass on feeding sites. Forage intake rate also increased linearly with plant quality. Male and female kiangs had similar intake rates. Likewise, gravid and lactating females had similar intake rates as barren and non-lactating females. At a coarse scale, kiangs spent longer time feeding in mesic than in xeric habitats, and spent more time feeding in early summer and fall than in late summer. Groups of adults with foals spent less time feeding than male groups and groups of adults without foals. Our findings suggest that kiangs use flexible foraging behaviours in relation to seasonal variations of vegetation quality and abundance, a likely outcome of the extreme seasonal conditions encountered on the Tibetan Plateau.