Allelopathy and plant invasions: traditional,congeneric, and bio-geographical approaches

A relatively small subset of exotic plant species competitively exclude their neighbors in invaded “recipient” communities but coexist with neighbors in their native habitat. Allelopathy has been argued as one of the mechanisms by which such exotics may become successful invaders. Three approaches have been used to examine allelopathy as a mechanism for invasion. The traditional approach examines exotic invasives in the same way that other native plants also suspected of allelopathic activities are studied. In this approach dose, fate, and replenishment of chemicals can provide powerful evidence for allelopathic processes. The bio-geographical approach often does not provide as much mechanistic evidence for allelopathy, but comparing the allelopathic effects of exotic invasives on species from their native and invaded communities yields stronger evidence than the traditional approach for whether or not allelopathy actually contributes to invasive success. The congeneric, or phylogenetic, approach involves comparative studies of exotic species with natives in the same genus or that are as closely related as possible. Congeneric approaches are limited in inference and have been used to study the role of natural enemies in exotic invasion, but this approach has not been widely used to study allelopathy and invasion. We discuss these three approaches and present a data set for congeneric Lantana and Prosopis to illustrate how the congeneric approach can be used, and use Centaurea maculosa and (±)-catechin to demonstrate experimentally how traditional and bio-geographic approaches can be integrated to shed light on allelopathy in exotic plant invasions.     

Plant litter effects on soil nutrient availability and vegetation dynamics: changes that occur when annual grasses invade shrub-steppe communities

Changes in the quantity and quality of plant litter occur in many ecosystems as they are invaded by exotic species, which impact soil nutrient cycling and plant community composition. Such changes in sagebrush-steppe communities are occurring with invasion of annual grasses (AG) into a perennial grass (PG) dominated system. We conducted a 5-year litter manipulation study located in the northern Great Basin, USA. Springtime litter was partially or completely removed in three communities with differing levels of invasion (invaded, mixed, and native) to determine how litter removal and litter biomass affected plant-available soil N and plant community composition. Litter biomass (prior to the removal treatment) was negatively correlated with plant-available N in the invaded community, but was positively correlated in the native community. Plant-available N had greater intra- and inter-annual fluctuations in the invaded compared to the mixed or native communities, but was not generally affected by removal treatments. Litter removal had negative effects on AG cover during a warm/dry year and negative effects on PG cover during a cool/wet year in the mixed community. Overall, the effectiveness of springtime litter manipulations on plant-available N were limited and weather dependent, and only removal treatments >75 % had effects on the plant community. Our study demonstrates how communities invaded by AGs have significantly increased temporal variability in nutrient cycling, which may decrease ecosystem stability. Further, we found that the ecological impacts from litter manipulation on sagebrush communities were dependent on the extent of AG invasion, the timing of removal, and seasonal precipitation.     

The Effects of Exotic Grasses on Litter Decomposition in a Hawaiian Woodland: The Importance of Indirect Effects

Exotic grasses and grass-fueled fires have altered plant species composition in the seasonal submontane woodlands of Hawaii Volcanoes National Park. These changes have altered both structural and functional aspects of the plant community, which could, in turn, have consequences for litter decomposition and nitrogen (N) dynamics. In grass-invaded unburned woodland, grass removal plots within the woodland, and woodland converted to grassland by fire, we compared whole-system fluxes and the contributions of individual species to annual aboveground fine litterfall and litterfall N, and litter mass and net N loss. We assessed the direct contribution of grass biomass to decomposition and N dynamics, and we determined how grasses affected decomposition processes indirectly via effects on native species and alteration of the litter layer microenvironment. Grasses contributed 35% of the total annual aboveground fine litterfall in the invaded woodland. However, total litterfall mass and N were not different between the invaded woodland and the grass removal treatment because of compensation by the native tree Metrosideros polymorpha, which increased litter production by 37% ± 5% when grasses were removed. The 0.3 g N m^–2/y^–1 contained in this production increase was equal to the N contained in grass litter. Litter production and litterfall N was lowest in the grassland due to the loss of native litter inputs. Decomposition of litterfall on an area basis was highest in the grass-invaded woodland. We attributed this effect to increased inherent decomposability of native litter in the presence of grasses because (a) the microenvironment of the three vegetation treatments had little effect on decomposition of common litter types and (b) M. polymorpha litter produced in the invaded woodland decomposed faster than that produced in the grass removal plots due to higher lignin concentrations in the latter than in the former. Area-weighted decomposition was lowest in the grassland due to the absence of native litter inputs. Across all treatments, most litter types immobilized N throughout the incubation, and litter net N loss on an area basis was not different among treatments. Our results support the idea that the effects of a plant species or growth form on decomposition cannot be determined in isolation from the rest of the community or from the direct effects of litter quality and quantity alone. In this dry woodland, exotic grasses significantly altered decomposition processes through indirect effects on the quantity and quality of litter produced by native species.     

Current plant diversity but not its soil legacy influences exotic plant invasion

植物多样性而非其土壤遗留效应影响外来植物入侵 植物多样性可以影响外来植物入侵,然而植物多样性的土壤遗留效应是否能够影响外来植物入侵目前仍不清楚。植物多样性能够改变土壤微生物群落和土壤理化性质,这种遗留效应可能会对该土壤中外来植物的生长产生影响。因此,我们假设植物多样性的土壤遗留效应会影响外来植物的入侵。为了检验该假说,我们开展了一个两阶段的植物-土壤反馈实验。在土壤驯化阶段,我们将12个植物物种(4种禾草植物、3种豆科植物和5种杂类草植物)分别单独种植,或者随机选择8个物种(包含3个功能型)混合种植在土壤中。在反馈阶段,我们将入侵植物三叶鬼针草(Bidens pilosa)分别与本地禾草荩草(Arthraxon hispidus)、本地杂类草翅果菊(Pterocypsela indica)或者同时与荩草和翅果菊种植在被驯化过的土壤中。研 究结果显示,三叶鬼针草相对于其本地竞争植物的生长取决于驯化植物和竞争植物物种的功能型。驯化植物的多样性对三叶鬼针草与其本地竞争植物之间的生长差异没有显著影响。然而,随着本地竞争植物物种多样性的增加,三叶鬼针草相对于其本地竞争植物的生长显著降低。这些结果表明,当前的植物多样性可以通过增加入侵植物和本地植物之间的生长不平衡性来减少外来植物的入侵。但是,植物多样性的土壤遗留效应对外来植物入侵的影响可能很小。     

The effect of soil legacy on competition and invasion by Acacia dealbata Link

Plant–soil feedbacks can exacerbate competition between invasive and native species, although the net effect of the interaction between soil biota and competition is likely to be species-specific. Very few studies have addressed the combined effect of soil and competition on plant performance and invasion by exotic woody species. This study explores plant growth and competition between Acacia dealbata and Pinus pinaster in three different soils—native, disturbed and invaded—in Portugal. The invasion of native P. pinaster forests by A. dealbata can be explained by the stronger competition ability of the exotic tree species. Competition is stronger in the native soil, allowing the establishment of A. dealbata in this soil and the displacement of P. pinaster. During invasion, A. dealbata changes soil conditions and establishes positive plant–soil feedbacks that promote its own germination and growth and increase P. pinaster mortality. Soil disturbance by the introduction of a different exotic species, Eucalyptus globulus, did not promote invasion by A. dealbata. We found a significant effect of soil legacy on both growth and competitive ability of the invasive A. dealbata. The ability of A. dealbata to outcompete the native P. pinaster in its own soil and the positive plant–soil feedbacks established after invasion are important mechanisms for A. dealbata invasion.     

Manipulating disturbance regimes and seeding to restore mesic Mediterranean grasslands

Question: Can managing disturbance regimes alone or in combination with seeding native species serve to shift the balance from exotic towards native species? Location: Central coast of California, USA. Methods: We measured vegetation composition for 10 yr in a manipulative experiment replicated at three sites. Treatments included no disturbance, grazing and clipping at three frequencies with and without litter removal. We seeded eight native species into clipped plots and compared cover in comparable plots with no seeding. Results: Regardless of frequency, clipping generally shifted community dominance from exotic annual grasses to exotic annual forbs, rather than consistently favoring native species. At one site, perennial grass cover decreased in no‐disturbance plots, but only after 4 yr. Litter removal had minimal impact on litter depth and plant community composition. Grazing had a highly variable effect on the abundance of different plant guilds across sites and years. Seeding increased abundance of only two of eight native species. Conclusions: Managing disturbance regimes alone is insufficient to restore native species guilds in highly‐invaded grasslands and seeding native species has highly variable success.     

Legacy effects overwhelm the short-term effects of exotic plant invasion and restoration on soil microbial community structure,enzyme activities,and nitrogen cycling

Plant invasions can have substantial consequences for the soil ecosystem, altering microbial community structure and nutrient cycling. However, relatively little is known about what drives these changes, making it difficult to predict the effects of future invasions. In addition, because most studies compare soils from uninvaded areas to long-established dense invasions, little is known about the temporal dependence of invasion impacts. We experimentally manipulated forest understory vegetation in replicated sites dominated either by exotic Japanese barberry (Berberis thunbergii), native Viburnums, or native Vacciniums, so that each vegetation type was present in each site-type. We compared the short-term effect of vegetation changes to the lingering legacy effects of the previous vegetation type by measuring soil microbial community structure (phospholipid fatty acids) and function (extracellular enzymes and nitrogen mineralization). We also replaced the aboveground litter in half of each plot with an inert substitute to determine if changes in the soil microbial community were driven by aboveground or belowground plant inputs. We found that after 2 years, the microbial community structure and function was largely determined by the legacy effect of the previous vegetation type, and was not affected by the current vegetation. Aboveground litter removal had only weak effects, suggesting that changes in the soil microbial community and nutrient cycling were driven largely by belowground processes. These results suggest that changes in the soil following either invasion or restoration do not occur quickly, but rather exhibit long-lasting legacy effects from previous belowground plant inputs.     

Plant–soil feedback during biological invasions: effect of litter decomposition from an invasive plant (Sphagneticola trilobata) on its native congener (S. calendulacea)

生物入侵过程中的植物-土壤反馈：一种入侵植物的凋落物分解对其本地近缘植物的影响 植物入侵可通过正或负的植物-土壤反馈效应改变土壤的生物和非生物性质,从而影响入侵栖息地的土壤理化性质。许多入侵物种的凋落物分解可增加土壤养分,降低本地植物多样性,并导致进一步的植物入侵。关于入侵植物凋落物在不同土壤类型及深度分解及反馈效应的研究依然很少。本研究旨在明确入侵植物南美蟛蜞菊(Sphagneticola trilobata)凋落物在不同土壤类型和不同土壤深度条件下的分解情 况及其对本地近缘植物蟛蜞菊(S. calendulacea)生理生长的影响。将装有南美蟛蜞菊凋落物的尼龙袋加入到不同深度(即0、2、4 和6 cm)的砂土、营养土和粘土中,经6个月的分解后,回收凋落物袋并计算分解速率,随后在凋落物分解处理后的土壤中种植本地蟛蜞菊,并在生长期结束时测量其生理生态指标。研究结果表明,所有处理土壤类型中,凋落物在土壤深度为2和4 cm处分解后显著增加了土壤养分,而对本 地蟛蜞菊的叶片叶绿素、叶氮含量等生长指标表现为负效应。因此,入侵植物南美蟛蜞菊凋落物分解对土壤养分表现为正的反馈效应,而对本地植物蟛蜞菊的生长表现为负效应。我们的研究结果还表明,入侵植物的凋落物分解对土壤和本地物种的影响还因凋落物分解所在的土壤深度而显著不同。未来的研究应侧重于入侵栖息地中更多本地和入侵物种的植物-土壤反馈效应,以及更多土壤类型和土壤深度的入侵植物凋落物效应。     

Exotic species as modifiers of ecosystem processes: Litter decomposition in native and invaded secondary forests of NW Argentina

Invasions of exotic tree species can cause profound changes in community composition and structure, and may even cause legacy effect on nutrient cycling via litter production. In this study, we compared leaf litter decomposition of two invasive exotic trees (Ligustrum lucidum and Morus sp.) and two dominant native trees (Cinnamomum porphyria and Cupania vernalis) in native and invaded (Ligustrum-dominated) forest stands in NW Argentina. We measured leaf attributes and environmental characteristics in invaded and native stands to isolate the effects of litter quality and habitat characteristics. Species differed in their decomposition rates and, as predicted by the different species colonization status (pioneer vs. late successional), exotic species decayed more rapidly than native ones. Invasion by L. lucidum modified environmental attributes by reducing soil humidity. Decomposition constants (k) tended to be slightly lower (−5%) for all species in invaded stands. High SLA, low tensile strength, and low C:N of Morus sp. distinguish this species from the native ones and explain its higher decomposition rate. Contrary to our expectations, L. lucidum leaf attributes were similar to those of native species. Decomposition rates also differed between the two exotic species (35% higher in Morus sp.), presumably due to leaf attributes and colonization status. Given the high decomposition rate of L. lucidum litter (more than 6 times that of natives) we expect an acceleration of nutrient circulation at ecosystem level in Ligustrum-dominated stands. This may occur in spite of the modified environmental conditions that are associated with L. lucidum invasion.     

Exotic plant invasion alters chaparral ecosystem resistance and resilience pre- and post-wildfire

Perturbations such as wildfire and exotic plant invasion have significant impacts on soils, and the extent to which invaded soils are resistant or resilient to these disturbances varies by ecosystem type. Replacement of shrublands by herbaceous exotics pre- and post-wildfire may drastically alter soil chemical and biological properties for an unknown duration. We assessed above and belowground resistance and resilience to exotic plant invasion both before and after a chaparral wildfire. We hypothesized that exotic plant species would change chemical characteristics of chaparral soils by altering litter and microbial inputs, and that controlling exotics and seeding native species would restore chemical characteristics to pre-invaded conditions. We additionally hypothesized that exotic plant species would slow succession above- and belowground, as well as recovery of post-wildfire chaparral structure and function. Plant species composition and soil nutrient pools and cycling rates were evaluated in mature and invaded chaparral pre- and post-wildfire. Exotic plant species were weeded and native species were seeded to assess impacts of exotic competition on native species recovery. Invasion did not impact all soil characteristics before fire, but increased soil C/N ratio, pH, and N cycling rates, and reduced NO₃-N availability. After fire, invasives slowed succession above- and belowground. Removal of exotics and seeding natives facilitated succession and resulted in plant composition similar to uninvaded, post-wildfire chaparral. The chaparral ecosystem was not resistant to impacts of invasion as indicated by altered soil chemistry and C and N cycling rates; however, short-term restoration led to recovery of extractable nitrogen availability indicating resilience of chaparral soils. This suggests that the permanence of exotic plant species, once established, represents a greater ecological challenge than exotic plant impacts on soils.     

Plant–soil feedbacks of exotic plant species across life forms: a meta-analysis

Invasive exotic plant species effects on soil biota and processes in their new range can promote or counteract invasions via changed plant–soil feedback interactions to themselves or to native plant species. Recent meta-analyses reveale that soil influenced by native and exotic plant species is affecting growth and performance of natives more strongly than exotics. However, the question is how uniform these responses are across contrasting life forms. Here, we test the hypothesis that life form matters for effects on soil and plant–soil feedback. In a meta-analysis we show that exotics enhanced C cycling, numbers of meso-invertebrates and nematodes, while having variable effects on other soil biota and processes. Plant effects on soil biota and processes were not dependent on life form, but patterns in feedback effects of natives and exotics were dependent on life form. Native grasses and forbs caused changes in soil that subsequently negatively affected their biomass, whereas native trees caused changes in soil that subsequently positively affected their biomass. Most exotics had neutral feedback effects, although exotic forbs had positive feedback effects. Effects of exotics on natives differed among plant life forms. Native trees were inhibited in soils conditioned by exotics, whereas native grasses were positively influenced in soil conditioned by exotics. We conclude that plant life form matters when comparing plant–soil feedback effects both within and between natives and exotics. We propose that impact analyses of exotic plant species on the performance of native plant species can be improved by comparing responses within plant life form.     

Interactive effects of global warming and ‘global worming’ on the initial establishment of native and exotic herbaceous plant species

The spread of exotic earthworms (‘worming’) and rising temperatures are expected to alter the biological, chemical and physical properties of many ecosystems, yet little is known about their potential interactive effects. We performed a laboratory microcosm experiment to investigate the effects of earthworms (anecic, endogeic, epigeic, or all three together) and 4°C warming on soil water content, litter turnover and seedling establishment of four native and four exotic herbaceous plant species. Warming and worming exerted independent as well as interactive effects on soil processes and plant dynamics. Warming reduced the water content of the upper soil layer, but only in the presence of earthworms. Litter removal increased in the presence of earthworms, the effect being most pronounced in the presence of anecic earthworms at ambient temperature. Exotic plant species were most influenced by earthworms (lower seedling number but higher biomass), whereas natives were most sensitive to warming (higher seedling number). This differential response resulted in significant interaction effects of earthworms and warming on abundance and richness of native relative to exotic plants as well as related shifts in plant species composition. Structural equation modeling allowed us to address possible mechanisms: direct effects of earthworms primarily affected exotic plants, whereas earthworms and warming indirectly and differentially affected native and exotic plants through changes in soil water content and surface litter. Invasive earthworms and warming are likely to interactively impact abiotic and biotic ecosystem properties. The invasion of epigeic and anecic species could select for plant species able to germinate on bare soil and tolerate drought, with the latter becoming more important in a warmer world. Thus earthworm invasion may result in simplified plant communities of increased susceptibility to the invasion of exotic plants.     

Do Native Parasitic Plants Cause More Damage to Exotic Invasive Hosts Than Native Non-Invasive Hosts? An Implication for Biocontrol

Field studies have shown that native, parasitic plants grow vigorously on invasive plants and can cause more damage to invasive plants than native plants. However, no empirical test has been conducted and the mechanism is still unknown. We conducted a completely randomized greenhouse experiment using 3 congeneric pairs of exotic, invasive and native, non-invasive herbaceous plant species to quantify the damage caused by parasitic plants to hosts and its correlation with the hosts' growth rate and resource use efficiency. The biomass of the parasitic plants on exotic, invasive hosts was significantly higher than on congeneric native, non-invasive hosts. Parasites caused more damage to exotic, invasive hosts than to congeneric, native, non-invasive hosts. The damage caused by parasites to hosts was significantly positively correlated with the biomass of parasitic plants. The damage of parasites to hosts was significantly positively correlated with the relative growth rate and the resource use efficiency of its host plants. It may be the mechanism by which parasitic plants grow more vigorously on invasive hosts and cause more damage to exotic, invasive hosts than to native, non-invasive hosts. These results suggest a potential biological control effect of native, parasitic plants on invasive species by reducing the dominance of invasive species in the invaded community.     

Does relatedness of natives used for soil conditioning influence plant-soil feedback of exotics?

The naturalisation hypothesis has been gaining attention recently as a possible mechanism to explain variations in invasion success. It predicts that exotic genera with native representatives should be less successful because of an overlap in resource use and of the existence of common specialised enemies. In this study, we tested whether native congenerics have more negative impact on exotic species than heterogenerics by increasing the effects of soil pathogens. We sampled soil in populations of three exotic species (Epilobium ciliatum, Impatiens parviflora and Stenactis annua) at sites with and without respective congeneric species. This soil was used as an inoculum for cultivating the first plant cohort, which included exotics, as well as native congenerics and heterogenerics. The conditioned soil was subsequently used for cultivating the second cohort of plants (exotics only). We found no consistent impact of relatedness of conditioning species on exotic growth. Although soil conditioned by congeneric E. hirsutum had the largest reduction on the performance of E. ciliatum, the final biomass of S. annua was lowest when grown in soil conditioned by itself. There was no effect of stimulating species on the biomass of I. parviflora. In both experimental phases, performance of exotics was improved when cultivated with sterilised inoculua, indicating the dominance of soil generalist pathogens. However, the biomass of S. annua was increased most by congeneric-stimulated inoculum from congeneric sites, suggesting a possible role for specialised symbionts. Our results suggest that variations in invasion success of at least some exotics may be affected by species-specific interactions mediated by the soil biota.     

Effects of exotic annual grass litter and local environmental gradients on annual plant community structure

Exotic annual grasses have been introduced into many semi-arid ecosystems worldwide, often to the detriment of native plant communities. The accumulation of litter from these grasses (i.e. residual dry biomass) has been demonstrated to negatively impact native plant communities and promote positive feedbacks to exotic grass persistence. More targeted experiments are needed, however, to determine the relative impact of exotic grass litter on plant community structure across local environmental gradients. We experimentally added exotic grass litter to annual forb-dominated open woodland communities positioned along natural canopy cover gradients in southwest Western Australia. These communities are an important component of this region’s plant biodiversity hotspot and are documented to be under threat from exotic annual grasses. After a one-year treatment period, we measured the effects of exotic grass litter, soil properties, and canopy cover on native and exotic species richness and abundance, as well as common species’ biomass and abundances. Plant community structure was more strongly influenced by soil properties and canopy cover than by grass litter. Total plant abundances per plot, however, were significantly lower in litter addition plots than control plots, a trend driven by native species. Exotic grass litter was also associated with lower abundances of one very common native species: Waitzia acuminata. Our results suggest that exotic grass litter limits the establishment of some native species in this system. Over multiple years, these subtle impacts may contribute substantially to the successful advancement of exotic species into this system, particularly in certain microenvironments.     

外来植物入侵对陆地生态系统地下碳循环及碳库的影响

生物入侵是当今全球性重大环境问题之一, 是全球变化的主要研究内容.评价外来植物入侵对于生态系统影响的研究多集中在地上部分,对于生态系统地下部分影响的研究相对较少.陆地生态系统地下部分对于生态系统过程的重要性之一体现在它处于生态系统碳分配过程的核心环节.入侵种通过影响群落凋落物的输入数量、质量以及输入时间,影响到对于土壤的碳输入,而入侵种与土著种根系的差异以及入侵种对微生物群落的影响是造成土壤呼吸强度发生变化的主要因素,前者土壤呼吸强度一般比后者高.多数研究表明外来植物入侵对生态系统地下碳循环和碳库产生影响,但由于入侵植物种类较多以及研究地点环境条件的不同,关于外来植物入侵对于土壤碳库和土壤有机碳矿化影响的研究结论并不统一.最后,提出了今后该研究领域应加强的一些建议和方向.     

The invasive species Alliaria petiolata (garlic mustard) increases soil nutrient availability in northern hardwood-conifer forests

The invasion of non-native plants can alter the diversity and activity of soil microorganisms and nutrient cycling within forests. We used field studies to analyze the impact of a successful invasive groundcover, Alliaria petiolata, on fungal diversity, soil nutrient availability, and pH in five northeastern US forests. We also used laboratory and greenhouse experiments to test three mechanisms by which A. petiolata may alter soil processes: (1) the release of volatile, cyanogenic glucosides from plant tissue; (2) the exudation of plant secondary compounds from roots; and (3) the decomposition of litter. Fungal community composition was significantly different between invaded and uninvaded soils at one site. Compared to uninvaded plots, plots invaded by A. petiolata were consistently and significantly higher in N, P, Ca and Mg availability, and soil pH. In the laboratory, the release of volatile compounds from the leaves of A. petiolata did not significantly alter soil N availability. Similarly, in the greenhouse, the colonization of native soils by A. petiolata roots did not alter soil nutrient cycling, implying that the exudation of secondary compounds has little effect on soil processes. In a leaf litter decomposition experiment, however, green rosette leaves of A. petiolata significantly increased the rate of decomposition of native tree species. The accelerated decomposition of leaf litter from native trees in the presence of A. petiolata rosette leaves shows that the death of these high-nutrient-content leaves stimulates decomposition to a greater extent than any negative effect that secondary compounds may have on the activity of the microbes decomposing the native litter. The results presented here, integrated with recent related studies, suggest that this invasive plant may change soil nutrient availability in such a way as to create a positive feedback between site occupancy and continued proliferation.     

Non-Additive Effects on Decomposition from Mixing Litter of the Invasive Mikania micrantha H.B.K. with Native Plants

A common hypothesis to explain the effect of litter mixing is based on the difference in litter N content between mixed species. Although many studies have shown that litter of invasive non-native plants typically has higher N content than that of native plants in the communities they invade, there has been surprisingly little study of mixing effects during plant invasions. We address this question in south China where Mikania micrantha H.B.K., a non-native vine, with high litter N content, has invaded many forested ecosystems. We were specifically interested in whether this invader accelerated decomposition and how the strength of the litter mixing effect changes with the degree of invasion and over time during litter decomposition. Using litterbags, we evaluated the effect of mixing litter of M. micrantha with the litter of 7 native resident plants, at 3 ratios: M₁ (1∶4, = exotic:native litter), M₂ (1∶1) and M₃ (4∶1, = exotic:native litter) over three incubation periods. We compared mixed litter with unmixed litter of the native species to identify if a non-additive effect of mixing litter existed. We found that there were positive significant non-additive effects of litter mixing on both mass loss and nutrient release. These effects changed with native species identity, mixture ratio and decay times. Overall the greatest accelerations of mixture decay and N release tended to be in the highest degree of invasion (mix ratio M₃) and during the middle and final measured stages of decomposition. Contrary to expectations, the initial difference in litter N did not explain species differences in the effect of mixing but overall it appears that invasion by M. micrantha is accelerating the decomposition of native species litter. This effect on a fundamental ecosystem process could contribute to higher rates of nutrient turnover in invaded ecosystems.     

Plant litter chemistry alters the content and composition of organic carbon associated with soil mineral and aggregate fractions in invaded ecosystems

Through the input of disproportionate quantities of chemically distinct litter, invasive plants may potentially influence the fate of organic matter associated with soil mineral and aggregate fractions in some of the ecosystems they invade. Although context dependent, these native ecosystems subjected to prolonged invasion by exotic plants may be instrumental in distinguishing the role of plant–microbe–mineral interactions from the broader edaphic and climatic influences on the formation of soil organic matter (SOM). We hypothesized that the soils subjected to prolonged invasion by an exotic plant that input recalcitrant litter (Japanese knotweed, Polygonum cuspidatum) would have a greater proportion of plant‐derived carbon (C) in the aggregate fractions, as compared with that in adjacent soil inhabited by native vegetation that input labile litter, whereas the soils under an invader that input labile litter (kudzu, Pueraria lobata) would have a greater proportion of microbial‐derived C in the silt‐clay fraction, as compared with that in adjacent soils that receive recalcitrant litter. At the knotweed site, the higher C content in soils under P. cuspidatum, compared with noninvaded soils inhabited by grasses and forbs, was limited to the macroaggregate fraction, which was abundant in plant biomarkers. The noninvaded soils at this site had a higher abundance of lignins in mineral and microaggregate fractions and suberin in the macroaggregate fraction, partly because of the greater root density of the native species, which might have had an overriding influence on the chemistry of the above‐ground litter input. At the kudzu site, soils under P. lobata had lower C content across all size fractions at a 0–5 cm soil depth despite receiving similar amounts of Pinus litter. Contrary to our prediction, the noninvaded soils receiving recalcitrant Pinus litter had a similar abundance of plant biomarkers across both mineral and aggregate fractions, potentially because of the higher surface area of soil minerals at this site. The plant biomarkers were lower in the aggregate fractions of the P. lobata‐invaded soils, compared with noninvaded pine stands, potentially suggesting a microbial co‐metabolism of pine‐derived compounds. These results highlight the complex interactions among litter chemistry, soil biota, and minerals in mediating soil C storage in unmanaged ecosystems; these interactions are particularly important under global changes that may alter plant species composition and hence the quantity and chemistry of litter inputs in terrestrial ecosystems.     

Fire and Litter Effects on Seedling Establishment in Western Oregon Upland Prairies

Prescribed burning is an important tool for managing and restoring prairies and other ecosystems. One effect of fire is plant litter removal, which can influence seedling establishment. Four experimental treatments (burned, clipped and raked to remove litter, burned with litter reapplied, and unmanipulated) were applied to 2 × 2.5–m plots in three western Oregon, United States, upland prairies to determine how burning affects seedling establishment. Seeds of common exotic and native prairie species were sowed into the experimental plots after treatments. Seedlings were censused the following spring. The experiment was repeated on each of the three sites, representing three common types of prairie vegetation: an Annual Exotic Grass site, a Perennial Exotic Grass site, and a Native Bunchgrass site. In both the Annual Exotic Grass and the Perennial Exotic Grass sites, burning significantly improved native, but not exotic, seedling establishment over those on unburned plots. Litter removal was a significant component of this burn effect, particularly on the Perennial Exotic Grass site. In these winter‐moist systems, the net effect of litter is to inhibit seedling establishment. Burning treatments on the Native Bunchgrass site significantly increased seedling establishment only of short‐lived exotic species. These results suggest that in prairie ecosystems similar to the Annual and Perennial Exotic Grass sites, prescribed burning followed by sowing native seeds can be an effective restoration technique. Burning alone or sowing alone would be counter‐productive, in the first case because increased establishment would come from exotic species and in the second case because establishment rates are low in unburned plots.