Soil Biota Facilitate Exotic Acer Invasions in Europe and North America

The primary hypothesis for successful exotic plant invasions is that the invaders have escaped the specialist consumers that control them (Enemy Release Hypothesis). However, few studies have rigorously tested this assertion with biogeographical experiments or considered the effects of soil organisms. We tested the Enemy Release Hypothesis and the enhanced role of mutualisms by comparing density patterns of the North American Acer negundo and European A. platanoides trees in their native and nonnative ranges. Invaders that have escaped their natural enemies are predicted to attain greater densities in nonnative than native ranges. To determine whether interactions with soil biota could explain the population distributions observed in the field, we compared the effects of sterilized and nonsterilized soil associated with Acer and non-Acer trees in native and nonnative ranges on the growth of seedlings in the greenhouse. In the field study, distances from focal trees to the nearest Acer conspecifics were 56– 77% less in their nonnative ranges than in their native ranges. In the greenhouse experiment, the effect of soil biota also differed between native and nonnative ranges of Acers. Relative to sterilized controls, soil associated with conspecifics and heterospecifics from the native ranges decreased the total biomass and relative change in height of Acer seedlings by 35% and 40%, respectively. Soil associated with conspecifics in the nonnative ranges decreased the biomass and relative change in height of Acer seedlings by 112% and 64%, respectively; but the soil associated with heterospecifics in the nonnative ranges increased biomass and relative change in height of Acer seedlings by 13% and 37%, respectively. Our results suggest that invasion of Acers is enhanced by soil biota associated with dominant native species and that the soil biota effect becomes more inhibitory as the Acers establish. The relative difference in soil biota effects between ranges supports the Enemy Release Hypothesis but also suggests that mutualists are relatively more beneficial to Acers in their nonnative ranges than in their native ranges. Mutualisms may be relatively more beneficial in nonnative ranges because the invader has escaped from the negative effect of natural enemies that may attenuate the positive effect of mutualists

Invasion Through Quantitative Effects: Intense Shade Drives Native Decline and Invasive Success

The effects of invasive nonnative species on community composition are well documented. However, few studies have determined the mechanisms by which invaders drive these changes. The literature indicates that many nonnative plant species alter light availability differently than natives in a given community, suggesting that shading may be such a mechanism. We compared light quantity (photosynthetically active radiation, PAR) and quality (red : far-red ratio, R:Fr) in riparian reaches heavily invaded by a nonnative tree (Acer platanoides) to that in an uninvaded forest and experimentally tested the effects of our measured differences in PAR and R:Fr on the survival, growth, and biomass allocation of seedlings of the dominant native species and Acer platanoides. Light conditions representative of the understory of Acer platanoides-invaded forest decreased survival of the native maple Acer glabrum by 28%; Amelanchier alnifolia by 32%; Betula occidentalis by 55%; Elymus glaucus by 46%; and Sorbus aucuparia by 52%, relative to seedlings growing in PAR similar to that of native understories. In contrast, Acer platanoides and the native shrub Symphoricarpos albus were not affected by reductions in PAR. Acer platanoides seedlings and saplings are uniquely adapted to shade relative to native species. Acer platanoides was the only species tested that decreased allocation to roots relative to shoots in the invaded forest vs. the native forest light conditions. Therefore it was the only species to demonstrate an adaptive response to the particular light environment associated with Acer platanoides invasion as predicted by optimal partitioning theory. The profound change in light quantity associated with Acer platanoides canopies appears to act as an important driver of native suppression and conspecific success in invaded riparian communities. Further research is necessary to determine whether the effect of nonnative plant-driven changes on light quantity and quality is a widespread mechanism negatively affecting resident species and facilitating invasion by nonnatives

Effects of Soil Biota from Different Ranges on Robinia Invasion: Acquiring Mutualists and Escaping Pathogens

The net effects of soil biota on exotic invaders can be variable, in part, because net effects are produced by many interacting mutualists and antagonists. Here we compared mutualistic and antagonistic biota in soils collected in the native, expanded, and invasive range of the black locust tree, Robinia pseudoacacia. Robinia formed nodules in all soils with a broad phylogenetic range of N-fixing bacteria, and leaf N did not differ among the different sources of soil. This suggests that the global expansion of Robinia was not limited by the lack of appropriate mutualistic N-fixers. Arbuscular mycorrhizal fungi (AMF) from the native range stimulated stronger positive feedbacks than AMF from the expanded or invasive ranges, a biogeographic difference not described previously for invasive plants. Pythium taxa collected from soil in the native range were not more pathogenic than those from other ranges; however, feedbacks produced by the total soil biota were more negative from soils from the native range than from the other ranges, overriding the effects of AMF. This suggests that escape from other pathogens in the soil or the net negative effects of the whole soil community may contribute to superior performance in invaded regions. Our results suggest that important regional evolutionary relationships may occur among plants and soil biota, and that net effects of soil biota may affect invasion, but in ways that are not easily explained by studying isolated components of the soil biota

Covariation in Plant Functional Traits and Soil Fertility within Two Species-Rich Forests

The distribution of plant species along environmental gradients is expected to be predictable based on organismal function. Plant functional trait research has shown that trait values generally vary predictably along broad-scale climatic and soil gradients. This work has also demonstrated that at any one point along these gradients there is a large amount of interspecific trait variation. The present research proposes that this variation may be explained by the local-scale sorting of traits along soil fertility and acidity axes. Specifically, we predicted that trait values associated with high resource acquisition and growth rates would be found on soils that are more fertile and less acidic. We tested the expected relationships at the species-level and quadrat-level (20×20 m) using two large forest plots in Panama and China that contain over 450 species combined. Predicted relationships between leaf area and wood density and soil fertility were supported in some instances, but the majority of the predicted relationships were rejected. Alternative resource axes, such as light gradients, therefore likely play a larger role in determining the interspecific variability in plant functional traits in the two forests studied

Effects of Mowing on Methane Uptake in a Semiarid Grassland in Northern China

Background: Mowing is a widely adopted management practice for the semiarid steppe in China and affects CH4 exchange. However, the magnitude and the underlying mechanisms for CH 4 uptake in response to mowing remain uncertain. Methodology/Principal Findings: In two consecutive growing seasons, we measured the effect of mowing on CH 4 uptake in a steppe community. Vegetation was mowed to 2 cm (M2), 5 cm (M5), 10 cm (M10), 15 cm (M15) above soil surface, respectively, and control was set as non-mowing (NM). Compared with control, CH4 uptake was substantially enhanced at almost all the mowing treatments except for M15 plots of 2009. CH4 uptake was significantly correlated with soil microbial biomass carbon, microbial biomass nitrogen, and soil moisture. Mowing affects CH 4 uptake primarily through its effect on some biotic factors, such as net primary productivity, soil microbial C\N supply and soil microbial activities, while soil temperature and moisture were less important. Conclusions/Significance: This study found that mowing affects the fluxes of CH4 in the semiarid temperate steppe of north China

Growth Response of Drought-Stressed Pinus sylvestris Seedlings to Single- and Multi-Species Inoculation with Ectomycorrhizal Fungi

Many trees species form symbiotic associations with ectomycorrhizal (ECM) fungi, which improve nutrient and water acquisition of their host. Until now it is unclear whether the species richness of ECM fungi is beneficial for tree seedling performance, be it during moist conditions or drought. We performed a pot experiment using Pinus sylvestris seedlings inoculated with four selected ECM fungi (Cenococcum geophilum, Paxillus involutus, Rhizopogon roseolus and Suillus granulatus) to investigate (i) whether these four ECM fungi, in monoculture or in species mixtures, affect growth of P. sylvestris seedlings, and (ii) whether this effect can be attributed to species number per se or to species identity. Two different watering regimes (moist vs. dry) were applied to examine the context-dependency of the results. Additionally, we assessed the activity of eight extracellular enzymes in the root tips. Shoot growth was enhanced in the presence of S. granulatus, but not by any other ECM fungal species. The positive effect of S. granulatus on shoot growth was more pronounced under moist (threefold increase) than under dry conditions (twofold increase), indicating that the investigated ECM fungi did not provide additional support during drought stress. The activity of secreted extracellular enzymes was higher in S. granulatus than in any other species. In conclusion, our findings suggest that ECM fungal species composition may affect seedling performance in terms of aboveground biomass

Effects of Tillage and Nitrogen Fertilizers on CH4 and CO2 Emissions and Soil Organic Carbon in Paddy Fields of Central China

Quantifying carbon (C) sequestration in paddy soils is necessary to help better understand the effect of agricultural practices on the C cycle. The objective of the present study was to assess the effects of tillage practices [conventional tillage (CT) and no-tillage (NT)] and the application of nitrogen (N) fertilizer (0 and 210 kg N ha−1) on fluxes of CH4 and CO2, and soil organic C (SOC) sequestration during the 2009 and 2010 rice growing seasons in central China. Application of N fertilizer significantly increased CH4 emissions by 13%–66% and SOC by 21%–94% irrespective of soil sampling depths, but had no effect on CO2 emissions in either year. Tillage significantly affected CH4 and CO2 emissions, where NT significantly decreased CH4 emissions by 10%–36% but increased CO2 emissions by 22%–40% in both years. The effects of tillage on the SOC varied with the depth of soil sampling. NT significantly increased the SOC by 7%–48% in the 0–5 cm layer compared with CT. However, there was no significant difference in the SOC between NT and CT across the entire 0–20 cm layer. Hence, our results suggest that the potential of SOC sequestration in NT paddy fields may be overestimated in central China if only surface soil samples are considered

Appendix A. Further details of methods used for measuring the effect of Acer platanoides on canopy structure and understory light and a table showing allometry of shrubs and saplings naturally occurring in shaded understories.

Further details of methods used for measuring the effect of Acer platanoides on canopy structure and understory light and a table showing allometry of shrubs and saplings naturally occurring in shaded understories

Supplement 1. Multirow spherical panoramas in QuickTime Virtual Reality (QTVR) format for depicting native riparian vegetation and riparian vegetation heavily invaded by Acer platanoides occurring adjacent to Rattlesnake Creek, Missoula, Montana, USA, in 2005.

<h2>File List</h2><blockquote> <p> <a href="native_riparian_forest.mov">native_riparian_forest.mov</a>  --  360º panorama in Quicktime Virtual Reality format </p> <p><a href="native_riparian_forest.jpg">native_riparian_forest.jpg</a>  --  still image in jpeg format </p> <p><a href="invaded_riparian_forest.mov">invaded_riparian_forest.mov</a>  --  360º panorama in Quicktime Virtual Reality format </p> <p><a href="invaded_riparian_forest.jpg">invaded_riparian_forest.jpg</a>  --  still image in jpeg format </p> </blockquote><h2>Description</h2><blockquote> <p> Multirow spherical (360°) panoramas in QuickTime Virtual Reality [QTVR] format are provided to depict native riparian vegetation and riparian vegetation heavily invaded by <i>Acer platanoides</i> occurring adjacent to Rattlesnake Creek, Missoula, Montana 2005. All of the movies are in QuickTime (*.mov) format. If you cannot view the movies, QuickTime is available at <a href="http://quicktime.apple.com">quicktime.apple.com</a>. Each QTVR movie provides the perspective of standing in an individual forest because the user is able to control the rotation of the field of view with their mouse. The same images can also be viewed in a much lower image quality jpg format. </p> </blockquote