Claudia Stein

Consequences of plant-soil feedbacks in invasion.

1. Plant species can influence soil biota, which in turn can influence the relative performance of plant species. These plant–soil feedbacks (PSFs) have been hypothesized to affect many community-level dynamics including species coexistence, dominance and invasion. 2. The importance of PSFs in exotic species invasion, although widely hypothesized, has been difficult to determine because invader establishment necessarily precedes invader-mediated PSFs. Here, we combine a spatial simulation model of invasion that incorporates PSFs with a meta-analysis that synthesizes published case studies describing feedbacks between pairs of native and exotic species. 3. While our spatial model confirmed the link between positive soil feedbacks (‘home’ advantage) for exotic species and exotic species spread, results were dependent on the initial abundance of the exotic species and the equivalence of dispersal and life history characteristics between exotic and native species. 4. The meta-analysis of 52 native–exotic pairwise feedback comparisons in 22 studies synthesized measures of native and exotic performance in soils conditioned by native and exotic species. The analysis indicated that the growth responses of native species were often greater in soil conditioned by native species than in soil conditioned exotic species (a ‘home’ advantage). The growth responses of exotic species were variable and not consistently related to species soil-conditioning effects. 5. Synthesis. Overlaying empirical estimates of pairwise PSFs with spatial simulations, we conclude that the empirically measured PSFs between native and exotic plant species are often not consistent with predictions of the spread of exotic species and mono-dominance. This is particularly the case when exotic species are initially rare and share similar dispersal and average fitness characteristics with native species. However, disturbance and other processes that increase the abundance of exotic species as well as the inclusion of species dispersal and life history differences can interact with PSF effects to explain the spread of invasive species

Specific bottom–up effects of arbuscular mycorrhizal fungi across a plant–herbivore–parasitoid system

The majority of plants are involved in symbioses with arbuscular mycorrhizal fungi (AMF), and these associations are known to have a strong influence on the performance of both plants and insect herbivores. Little is known about the impact of AMF on complex trophic chains, although such effects are conceivable. In a greenhouse study we examined the effects of two AMF species, Glomus intraradices and G. mosseae on trophic interactions between the grass Phleum pratense, the aphid Rhopalosiphum padi, and the parasitic wasp Aphidius rhopalosiphi. Inoculation with AMF in our study system generally enhanced plant biomass (+5.2%) and decreased aphid population growth (−47%), but there were no fungal species-specific effects. When plants were infested with G. intraradices, the rate of parasitism in aphids increased by 140% relative to the G. mosseae and control treatment. When plants were associated with AMF, the developmental time of the parasitoids decreased by 4.3% and weight at eclosion increased by 23.8%. There were no clear effects of AMF on the concentration of nitrogen and phosphorus in plant foliage. Our study demonstrates that the effects of AMF go beyond a simple amelioration of the plants’ nutritional status and involve rather more complex species-specific cascading effects of AMF in the food chain that have a strong impact not only on the performance of plants but also on higher trophic levels, such as herbivores and parasitoids.

Impact of invertebrate herbivory in grasslands depends on plant species diversity

Invertebrate herbivores are ubiquitous in most terrestrial ecosystems, and theory predicts that their impact on plant community biomass should depend on diversity and productivity of the associated plant communities. To elucidate general patterns in the relationship between invertebrate herbivory, plant diversity, and productivity, we carried out a long-term herbivore exclusion experiment at multiple grassland sites in a mountainous landscape of central Germany. Over a period of five years, we used above- and belowground insecticides as well as a molluscicide to manipulate invertebrate herbivory at 14 grassland sites, covering a wide range of plant species diversity (13-38 species/m2) and aboveground plant productivity (272-1125 g x m(-2) x yr(-1)), where plant species richness and productivity of the sites were not significantly correlated. Herbivore exclusion had significant effects on the plant communities: it decreased plant species richness and evenness, and it altered plant community composition. In particular, exclusion of belowground herbivores promoted grasses at the expense of herbs. In contrast to our expectation, herbivore effects on plant community biomass were not influenced by productivity. However, effect size of invertebrate herbivores was negatively correlated with plant diversity of the grasslands: the effect of herbivory on biomass tended to be negative at sites of high diversity and positive at sites of low diversity. In general, the effects of aboveground herbivores were relatively small as compared to belowground herbivores, which were important drivers of plant community composition. Our study is the first to show that variation in the effects of invertebrate herbivory on plant communities across a landscape is significantly influenced by plant species richness.

Staged invasions across disparate grasslands: effects of seed provenance, consumers and disturbance on productivity and species richness.

Exotic plant invasions are thought to alter productivity and species richness, yet these patterns are typically correlative. Few studies have experimentally invaded sites and asked how addition of novel species influences ecosystem function and community structure and examined the role of competitors and/or consumers in mediating these patterns. We invaded disturbed and undisturbed subplots in and out of rodent exclosures with seeds of native or exotic species in grasslands in Montana, California and Germany. Seed addition enhanced aboveground biomass and species richness compared with no-seeds-added controls, with exotics having disproportionate effects on productivity compared with natives. Disturbance enhanced the effects of seed addition on productivity and species richness, whereas rodents reduced productivity, but only in Germany and California. Our results demonstrate that experimental introduction of novel species can alter ecosystem function and community structure, but that local filters such as competition and herbivory influence the magnitude of these impacts.

Herbivore preference drives plant community composition

Herbivores are important drivers of plant species coexistence and community assembly. However, detailed mechanistic information on how herbivores affect dominance hierarchies between plant species is scarce. Here, we used data of a multi-site herbivore exclusion experiment in grasslands to assess changes in the cover of 28 plant species in response to aboveground pesticide. application. Moreover, we assessed species-specific values of plant defense of these 28 species measured as the performance of a generalist caterpillar, and the preference of the caterpillar and a slug species in no-choice and choice feeding experiments, respectively. We show that more preferred species in the feeding experiments were those that increased in cover after herbivore exclusion in the field, whereas less preferred ones decreased. Herbivore performance and several measured leaf traits were not related to the change in plant cover in the field in response to herbivore removal. Additionally, the generalist slug and the generalist caterpillar preferred and disliked the same plant species, indicating that they perceive the balance between defense and nutritional value similarly. We conclude that the growth-defense trade-off in grassland species acts via the preference of herbivores and that among-species variation in plant growth and preference to herbivores drives plant community composition.

Evaluating Ecosystem Services Provided by Non-Native Species: An Experimental Test in California Grasslands

The concept of ecosystem services – the benefits that nature provides to humans society – has gained increasing attention over the past decade. Increasing global abiotic and biotic change, including species invasions, is threatening the secure delivery of these ecosystem services. Efficient evaluation methods of ecosystem services are urgently needed to improve our ability to determine management strategies and restoration goals in face of these new emerging ecosystems. Considering a range of multiple ecosystem functions may be a useful way to determine such strategies. We tested this framework experimentally in California grasslands, where large shifts in species composition have occurred since the late 1700s. We compared a suite of ecosystem functions within one historic native and two non-native species assemblages under different grazing intensities to address how different species assemblages vary in provisioning, regulatory and supporting ecosystem services. Forage production was reduced in one non-native assemblage (medusahead). Cultural ecosystem services, such as native species diversity, were inherently lower in both non-native assemblages, whereas most other services were maintained across grazing intensities. All systems provided similar ecosystem services under the highest grazing intensity treatment, which simulated unsustainable grazing intensity. We suggest that applying a more comprehensive ecosystem framework that considers multiple ecosystem services to evaluate new emerging ecosystems is a valuable tool to determine management goals and how to intervene in a changing ecosystem.

Colour morph related performance in the meadow grasshopper Chorthippus parallelus (Orthoptera, Acrididae)

Abstract 1. Polymorphism has been described for a number of herbivorous insects, but little is known about whether differences in body colour cause fitness differences. In Chorthippus parallelus, three main colour morphs occur, namely brown, green, and dorsally striped.

Functional diversity increases ecological stability in a grazed grassland

Understanding the factors governing ecological stability in variable environments is a central focus of ecology. Functional diversity can stabilize ecosystem function over time if one group of species compensates for an environmentally driven decline in another. Although intuitively appealing, evidence for this pattern is mixed. We hypothesized that diverse functional responses to rainfall will increase the stability of vegetation cover and biomass across rainfall conditions, but that this effect depends on land-use legacies that maintain functional diversity. We experimentally manipulated grazing in a California grassland to create land-use legacies of low and moderate grazing, across which we implemented rainout shelters and irrigation to create dry and wet conditions over 3 years. We found that the stability of the vegetation cover was greatly elevated and the stability of the biomass was slightly elevated across rainfall conditions in areas with histories of moderate grazing. Initial functional diversity—both in the seed bank and aboveground—was also greater in areas that had been moderately grazed. Rainfall conditions in conjunction with this grazing legacy led to different functional diversity patterns over time. Wet conditions led to rapid declines in functional diversity and a convergence on resource-acquisitive traits. In contrast, consecutively dry conditions maintained but did not increase functional diversity over time. As a result, grazing practices and environmental conditions that decrease functional diversity may be associated with lasting effects on the response of ecosystem functions to drought. Our results demonstrate that theorized relationships between diversity and stability are applicable and important in the context of working grazed landscapes.

Grassland Arthropods Are Controlled by Direct and Indirect Interactions with Cattle but Are Largely Unaffected by Plant Provenance

Cattle grazing and invasion by non-native plant species are globally-ubiquitous changes occurring to plant communities that are likely to reverberate through whole food webs. We used a manipulative field experiment to quantify how arthropod community structure differed in native and non-native California grassland communities in the presence and absence of grazing. The arthropod community was strongly affected by cattle grazing: the biovolume of herbivorous arthropods was 79% higher in grazed than ungrazed plots, whereas the biovolume of predatory arthropods was 13% higher in ungrazed plots. In plots where non-native grasses were grazed, arthropod biovolume increased, possibly in response to increased plant productivity or increased nutritional quality of rapidly-growing annual plants. Grazing may thus affect plant biomass both through the direct removal of biomass, and through arthropod-mediated impacts. We also expected the arthropod community to differ between native and non-native plant communities; surprisingly, arthropod richness and diversity did not vary consistently between these grass community types, although arthropod abundance was slightly higher in plots with native and ungrazed grasses. These results suggest that whereas cattle grazing affects the arthropod community via direct and indirect pathways, arthropod community changes commonly associated with non-native plant invasions may not be due to the identity or dominance of the invasive species in those systems, but to accompanying changes in plant traits or functional group composition, not seen in this experiment because of the similarity of the plant communities.

Grazing weakens temporal stabilizing effects of diversity in the Eurasian steppe

Abstract Many biodiversity experiments have demonstrated that plant diversity can stabilize productivity in experimental grasslands. However, less is known about how diversity–stability relationships are mediated by grazing. Grazing is known for causing species losses, but its effects on plant functional groups (PFGs) composition and species asynchrony, which are closely correlated with ecosystem stability, remain unclear. We conducted a six‐year grazing experiment in a semi‐arid steppe, using seven levels of grazing intensity (0, 1.5, 3.0, 4.5, 6.0, 7.5, and 9.0 sheep per hectare) and two grazing systems (i.e., a traditional, continuous grazing system during the growing period (TGS), and a mixed one rotating grazing and mowing annually (MGS)), to examine the effects of grazing system and grazing intensity on the abundance and composition of PFGs and diversity–stability relationships. Ecosystem stability was similar between mixed and continuous grazing treatments. However, within the two grazing systems, stability was maintained through different pathways, that is, along with grazing intensity, persistence biomass variations in MGS, and compensatory interactions of PFGs in their biomass variations in TGS. Ecosystem temporal stability was not decreased by species loss but rather remain unchanged by the strong compensatory effects between PFGs, or a higher grazing‐induced decrease in species asynchrony at higher diversity, and a higher grazing‐induced increase in the temporal variation of productivity in diverse communities. Ecosystem stability of aboveground net primary production was not related to species richness in both grazing systems. High grazing intensity weakened the temporal stabilizing effects of diversity in this semi‐arid grassland. Our results demonstrate that the productivity of dominant PFGs is more important than species richness for maximizing stability in this system. This study distinguishes grazing intensity and grazing system from diversity effects on the temporal stability, highlighting the need to better understand how grazing regulates ecosystem stability, plant diversity, and their synergic relationships.