Soil Nitrogen Conditions Approach Preinvasion Levels following Restoration of Nitrogen-Fixing Black Locust (Robinia pseudoacacia) Stands in a Pine–Oak Ecosystem

Restoring native plant communities on sites formerly occupied by invasive nitrogen‐fixing species poses unique problems due to elevated soil nitrogen availability. Mitigation practices that reduce available nitrogen may ameliorate this problem. We evaluated the effects of tree removal followed by soil preparation or mulching on native plant growth and soil nitrogen transformations in a pine–oak system formerly occupied by exotic nitrogen‐fixing Black locust (Robinia pseudoacacia) trees. Greenhouse growth experiments with native grasses, Andropogon gerardii and Sorghastrum nutans, showed elevated relative growth rates in soils from Black locust compared with pine–oak stands. Field soil nutrient concentrations and rates of net nitrification and total net N‐mineralization were compared 2 and 4 years since Black locust removal and in control sites. Although soil nitrogen concentrations and total net N‐mineralization rates in the restored sites were reduced to levels that were similar to paired pine–oak stands after only 2 years, net nitrification rates remained 3–34 times higher in the restored sites. Other nutrient ion concentrations (Ca, Mg) and organic matter content were reduced, whereas phosphorus levels remained elevated in restored sites. Thus, 2–4 years following Black locust tree removal and soil horizon mixing achieved through site preparation, the concentrations of many soil nutrients returned to preinvasion levels. However, net nitrification rates remained elevated; cover cropping or carbon addition during restoration of sites invaded by nitrogen fixers could increase nitrogen immobilization and/or reduce nitrate availability, making sites more amenable to native plant establishment.     

<Emphasis Type="Italic">Hedychium gardnerianum</Emphasis> Invasion into Hawaiian Montane Rainforest: Interactions Among Litter Quality,Decomposition Rate,and Soil Nitrogen Availability

Few studies have examined the invasion of understory species into closed-canopy forests and, despite inter-specific differences in litter quality and quantity between understory and dominant canopy trees, the influence of understory invasions on soil nitrogen (N) cycling remains unknown. This paper examines litter quality and decomposition of kahili ginger (Hedychium gardnerianum), an invasive understory herb, to determine the influence of this species on N cycling in a Hawaiian montane rainforest. To examine the potential feedback between increased soil N availability and litter decomposition, litter from the invasive ginger, a native tree, and native tree fern was collected from unfertilized and fertilized plots and decomposed in a reciprocal transplant design. Hedychium litter decomposed faster than litter from the two native species. Across species, decomposition rates were negatively correlated with litter lignin content. Despite rapid decomposition rates of Hedychium litter, soil nitrogen availability and rates of net mineralization in the soil were similar in invaded and uninvaded plots. Nitrogen cycling at this site may be more strongly influenced by native species, which contribute the most to overall stand biomass. A negative effect of fertilization on the decomposition of Hedychium litter suggests that a negative feedback between litter quality and soil N availability may exist over longer timescales.     

Does Eucalyptus grandis invasion and removal affect soils and vegetation in the Eastern Cape Province,South Africa?

Many invasive alien plants drive changes in native community composition, structure and diversity. They alter soil nutrient regimes of native communities and affect native plant recovery outcomes following their removal. We assessed whether Eucalyptus grandis invasion and removal alters the soil physico‐chemical properties and native vegetation recovery in the Eastern Cape Province, South Africa. We collected samples from topsoil in E. grandis invaded sites (canopy cover > 75%), cleared sites (eight years ago) and native sites (canopy cover > 80%) and quantified soil moisture, concentrations of soil macro elements (N, C and P), pH and exchangeable cations (K, Ca, Mg, Na) as well as measured soil water repellency using the Water Droplet Penetration Time and infiltration. We conducted vegetation surveys in plots measuring 10 × 10 m. Invasion by E. grandis had varying effects on soil physico‐chemical properties, causing increase in soil pH and P, while decreasing total N and C. The removal of E. grandis also showed varying effects on soil physico‐chemical properties, but seems to have further triggered the loss of some soil nutrients (especially soil P). Soil water repellency (a measure of soil compaction) has improved in cleared sites to non‐repellent soils compared to repellent soils in invaded site. Eucalyptus grandis reduced species richness of the invaded sites. The presence of native species on cleared sites indicates a positive trajectory towards vegetation recovery. We conclude that E. grandis invasion and removal trigger varying effects on soil properties (both increases and decreases). For soil and vegetation restoration of cleared sites to be effective, active restoration techniques such as soil transfer, nutrient manipulation and native plant seeding should be considered.     

Virulence of oomycete pathogens from Phragmites australis‐invaded and noninvaded soils to seedlings of wetland plant species

Soil pathogens affect plant community structure and function through negative plant–soil feedbacks that may contribute to the invasiveness of non‐native plant species. Our understanding of these pathogen‐induced soil feedbacks has relied largely on observations of the collective impact of the soil biota on plant populations, with few observations of accompanying changes in populations of specific soil pathogens and their impacts on invasive and noninvasive species. As a result, the roles of specific soil pathogens in plant invasions remain unknown. In this study, we examine the diversity and virulence of soil oomycete pathogens in freshwater wetland soils invaded by non‐native Phragmites australis (European common reed) to better understand the potential for soil pathogen communities to impact a range of native and non‐native species and influence invasiveness. We isolated oomycetes from four sites over a 2‐year period, collecting nearly 500 isolates belonging to 36 different species. These sites were dominated by species of Pythium, many of which decreased seedling survival of a range of native and invasive plants. Despite any clear host specialization, many of the Pythium species were differentially virulent to the native and non‐native plant species tested. Isolates from invaded and noninvaded soils were equally virulent to given individual plant species, and no apparent differences in susceptibility were observed between the collective groups of native and non‐native plant species.     

Invasion by <Emphasis Type="Italic">Aegilops triuncialis</Emphasis> (Barb Goatgrass) Slows Carbon and Nutrient Cycling in a Serpentine Grassland

Invasive plant species alter plant community composition and ecosystem function. In the United States, California native grasslands have been displaced almost completely by invasive annual grasses, with serpentine grasslands being one of the few remaining refugia for California grasslands. This study examined how the invasive annual grass, Aegilops triuncialis, has altered decomposition processes in a serpentine annual grassland. Our objectives were to (1) assess howA. triuncialis alters primary productivity and litter tissue chemistry, (2) determine whether A. triuncialis litter is more recalcitrant to decomposition than native litter, and (3) evaluate whether differences in the soil microbial community in A. triuncialis-invaded and native-dominated areas result in different decomposition rates of invasive and/or native plant litter. In invaded plant patches, A. triuncialis was approximately 50% of the total plant cover, in contrast to native plant patches in which A. triuncialis was not detected and native plants comprised over 90% of the total plant cover. End-of-season aboveground biomass was 2-fold higher in A. triuncialis dominated plots compared to native plots; however, there was no significant difference in belowground biomass. Both above- and below-ground plant litter from A. triuncialis plots had significantly higher lignin:N and C:N ratios and lower total N, P, and K than litter from native plant plots. Aboveground litter from native plots decomposed more rapidly than litter from A. triuncialis plots, although there was no difference in decomposition of belowground tissues. Soil microbial community composition associated with different soil patch types had no effect on decomposition rates. These data suggest that plant invasion impacts decomposition and nutrient cycling through changes in plant community tissue chemistry and biomass production.     

An invasive aster (Ageratina adenophora) invades and dominates forest understories in China: altered soil microbial communities facilitate the invader and inhibit natives

Exotic plant invasion may alter underground microbial communities, and invasion-induced changes of soil biota may also affect the interaction between invasive plants and resident native species. Increasing evidence suggests that feedback of soil biota to invasive and native plants leads to successful exotic plant invasion. To examine this possible underlying invasion mechanism, soil microbial communities were studied where Ageratina adenophora was invading a native forest community. The plant–soil biota feedback experiments were designed to assess the effect of invasion-induced changes of soil biota on plant growth, and interactions between A. adenophora and three native plant species. Soil analysis showed that nitrate nitrogen (NO₃⁻-N), ammonium nitrogen (NH₄⁺-N), and available P and K content were significantly higher in a heavily invaded site than in a newly invaded site. The structure of the soil microbial community was clearly different in all four sites. Ageratina adenophora invasion strongly increased the abundance of soil VAM (vesicular-arbuscular mycorrhizal fungi) and the fungi/bacteria ratio. A greenhouse experiment indicated that the soil biota in the heavily invaded site had a greater inhibitory effect on native plant species than on A. adenophora and that soil biota in the native plant site inhibited the growth of native plant species, but not of A. adenophora. Soil biota in all four sites increased A. adenophora relative dominance compared with each of the three native plant species and soil biota in the heavily invaded site had greater beneficial effects on A. adenophora relative dominance index (20% higher on average) than soil biota in the non-invaded site. Our results suggest that A. adenophora is more positively affected by the soil community associated with native communities than are resident natives, and once the invader becomes established it further alters the soil community in a way that favors itself and inhibits natives, helping to promote the invasion. Soil biota alteration after A. adenophora establishment may be an important part of its invasion process to facilitate itself and inhibit native plants.     

Forest Invasibility in Communities in Southeastern New York

While biological invasions have been the subject of considerable attention both historically and recently, the factors controlling the susceptibility of communities to plant invasions remain controversial. We surveyed 44 sites in southeastern New York State to examine the relationships between plant community characteristics, soil characteristics, and nonnative plant invasion. Soil nitrogen mineralization and nitrification rates were strongly related to the degree of site invasion (F= 30.2, P < 0.0001 and F= 11.8, P < 0.005, respectively), and leaf C : N ratios were negatively correlated with invasion (R ²= 0.22, P < 0.0001). More surprisingly, there was a strong positive relationship between soil calcium levels and the degree of site invasion (partial r= 0.70, P < 0.01), and there were also positive relationships between invasion and soil magnesium and phosphorus. We found, in addition, a positive factor-ceiling relationship between native species diversity and invasive species diversity. This positive relationship between native and invasive diversity contradicts earlier hypotheses concerning the relationships between species diversity and invasion, but supports some recent findings. Cluster analysis distinguished two broad forest community types at our sites: pine barrens and mixed hardwood communities. Invaders were significantly more abundant in mixed hardwood than in pine barrens communities (Mann–Whitney U = 682.5, P < 0.0001). Even when evaluating the mixed hardwood communities alone, invasion remained significantly positively correlated with soil fertility (calcium, magnesium, and net nitrogen mineralization rates). Soil texture and pH were not useful predictors of the degree to which forests were invaded. Nitrogen and calcium are critical components of plant development, and species better able to take advantage of increased nutrient availability may out-perform others at sites with higher nutrient levels. These results have implications for areas such as the eastern United States, where anthropogenic changes in the availability of nitrogen and calcium are affecting many plant communities.     

内生真菌对氮添加羽茅根际土壤特性和微生物群落的影响

内生真菌不仅能改变与其共生植物的生理和生长指标,还可通过宿主植物间接对土壤的理化性质和微生物群落结构产生影响。以天然禾草——羽茅(Achnatherum sibiricum)为研究材料,探究在不同施氮水平下,内生真菌的种类对不同基因型的宿主植物根际土壤理化性质和微生物群落产生何种影响。结果表明,内生真菌侵染显著提高了羽茅根际土壤的pH值和微生物总量,但降低了土壤中真菌与细菌的比值。同时,土壤的pH值还受到了内生真菌种类的影响,其中感染Epichlo3 sibirica的羽茅根际土壤pH显著高于感染Epichlo3 gansuense-1的羽茅,而感染Epichlo3 gansuense-2的羽茅根际土壤pH与感染E.gansuensis-1、E.sibirica菌的羽茅相比没有显著差异。另外,内生真菌感染与否、内生真菌种类、施氮量以及宿主植物基因型对土壤总碳、总氮、微生物及碳矿化能力均无显著影响。     

The invasive alien plant species Solidago gigantea alters ecosystem properties across habitats with differing fertility

Question: Invasive alien plants can affect biomass production and rates of biogeochemical cycling. Do the direction and intensity of such effects depend upon the functional traits of native and alien species and upon the properties of the invaded habitat, with the same alien species having differing impacts in different habitats? Location: Lowlands of Switzerland. Methods: Fourteen grassland and wetland sites invaded by Solidago gigantea and widely differing in biomass production and soil P availability were surveyed. To determine whether the impact of the species was related to site fertility, we compared the invaded and native vegetation in terms of biomass, species composition, plant traits and soil properties. Results: S. gigantea generally increased the above‐ground biomass production of the vegetation and soil C content, while reducing nutrient concentrations in biomass and N availability in the soil. However, it had no significant effect on plant species richness, soil respiration, soil pH and P availability. Leaves of S. gigantea had a greater C content than those of native species; other leaf traits and root phosphatase activity did not differ significantly. Conclusions: Our results suggest that a conservative nutrient‐use strategy allows S. gigantea to invade a broad range of habitats. The observed effects of invasion did not vary according to biomass production of the invaded sites, but some effects did depend on soil P availability, being more pronounced at more P‐rich sites. Thus, the full range of invaded habitats should be considered in studying the potential impact of plant invasions on ecosystem processes.     

Thyme travels: 15N isoscapes of Thymus vulgaris L. invasion in lightly grazed pastoral communities

Alterations to ecosystem nitrogen (N) cycling by introduced plant species may increase the invasibility of habitat providing a positive feedback for the introduced species to become invasive. Spatial patterns of foliar and soil δ¹⁵N ratios reflect variation in rates and process of N‐cycling across invaded landscapes and provide insight into N‐source uptake and utilization strategies of invasive plant species. To evaluate invasion‐associated changes in soil and foliar δ¹⁵N at different scales: regional (among different sites), local (between north‐ and south‐facing aspect at the same site), and microsite (within populations in the same community), we measured foliar and soil δ¹⁵ N, animal faeces cover (as a proxy for grazing intensity) and N₂‐fixing species cover from inside to outside Thymus vulgaris L. (thyme)‐invaded lightly grazed pastoral communities in Central Otago, southern South Island, New Zealand. Mean thyme foliar δ¹⁵N were near‐zero across the invaded landscape, and did not change across the advancing edge of invasion or with aspect. There was no evidence that associations with N₂‐fixing species provide a potential N source. Soil δ¹⁵N was lower inside of thyme compared to at the edge or outside of thyme and was varied between aspects at some sites. Animal faeces cover as a proxy for grazing intensity explained only 23% of this observed variation of soil δ¹⁵N. Thyme invasion may result in lowered soil δ¹⁵N reflecting alterations to N dynamics. Associated invasion‐related impacts of animal grazing may also impact soil δ¹⁵N. Further studies are required to distinguish the underlying mechanism responsible for the observed patterns of foliar and soil δ¹⁵N values across thyme‐invaded Central Otago landscapes.     

Woody Plant Encroachment by <Emphasis Type="Italic">Juniperus virginiana</Emphasis> in a Mesic Native Grassland Promotes Rapid Carbon and Nitrogen Accrual

The cover and abundance of Juniperus virginiana L. in the U.S. Central Plains are rapidly increasing, largely as a result of changing land-use practices that alter fire regimes in native grassland communities. Little is known about how conversion of native grasslands to Juniperus-dominated forests alters soil nutrient availability and ecosystem storage of carbon (C) and nitrogen (N), although such land-cover changes have important implications for local ecosystem dynamics, as well as regional C and N budgets. Four replicate native grasslands and adjacent areas of recent J. virginiana encroachment were selected to assess potential changes in soil N availability, leaf-level photosynthesis, and major ecosystem C and N pools. Net N mineralization rates were assessed in situ over two years, and changes in labile soil organic pools (potential C and N mineralization rates and microbial biomass C and N) were determined. Photosynthetic nitrogen use efficiencies (PNUE) were used to examine differences in instantaneous leaf-level N use in C uptake. Comparisons of ecosystem C and N stocks revealed significant C and N accrual in both plant biomass and soils in these newly established forests, without changes in labile soil N pools. There were few differences in monthly in situ net N mineralization rates, although cumulative annual net N mineralization was greater in forest soils compared to grasslands. Conversely, potential C mineralization was significantly reduced in forest soils. Encroachment by J. virginiana into grasslands results in rapid accretion of ecosystem C and N in plant and soil pools with little apparent change in N availability. Widespread increases in the cover of woody plants, like J. virginiana, in areas formerly dominated by graminoid species suggest an increasing role of expanding woodlands and forests as regional C sinks in the central U.S.     

The Influence of Soil Inoculum and Nitrogen Availability on Restoration of High-Elevation Steppe Communities Invaded by Bromus tectorum

Cheatgrass ( Bromus tectorum L.) is an exotic annual grass that has invaded approximately 40,000,000 ha of rangelands in the United States, including montane ecosystems that are important habitats for wildlife and livestock. In addition to well-understood mechanisms by which Cheatgrass gains competitive advantage, recent studies have shown that Cheatgrass may also change the associated soil microbial community to impact native perennial plants and promote the persistence of Cheatgrass. Furthermore, reducing plant-available N represents a tool for initiating conditions that accelerate successional change from annual- to perennial-dominated communities. At a montane, mixed shrub–grassland Cheatgrass-dominated site in Colorado, we applied sucrose to reduce available N, and we added soil from a native plant community in order to reestablish the microbial community. This approach tested the idea that intact native soil microbial communities may enhance the beneficial effect of reducing soil N availability in a restoration setting. By the end of the experiment, reduced N availability decreased Cheatgrass by 9.8%, non-native annual/biennial plant cover by 15.0%, and increased relative perennial plant cover by 13.4%; soil inoculation reduced Cheatgrass by 7.6% and increased perennial abundance by 11.3%. Soil inoculum additions and reduced N availability both contributed toward restoring a perennial-dominated community and demonstrates that addition of native soil inoculum may be a useful tool for restoration efforts.     

Soil pathogen communities associated with native and non‐native Phragmites australis populations in freshwater wetlands

Soil pathogens are believed to be major contributors to negative plant–soil feedbacks that regulate plant community dynamics and plant invasions. While the theoretical basis for pathogen regulation of plant communities is well established within the plant–soil feedback framework, direct experimental evidence for pathogen community responses to plants has been limited, often relying largely on indirect evidence based on above‐ground plant responses. As a result, specific soil pathogen responses accompanying above‐ground plant community dynamics are largely unknown. Here, we examine the oomycete pathogens in soils conditioned by established populations of native noninvasive and non‐native invasive haplotypes of Phragmites australis (European common reed). Our aim was to assess whether populations of invasive plants harbor unique communities of pathogens that differ from those associated with noninvasive populations and whether the distribution of taxa within these communities may help to explain invasive success. We compared the composition and abundance of pathogenic and saprobic oomycete species over a 2‐year period. Despite a diversity of oomycete taxa detected in soils from both native and non‐native populations, pathogen communities from both invaded and noninvaded soils were dominated by species of Pythium. Pathogen species that contributed the most to the differences observed between invaded and noninvaded soils were distributed between invaded and noninvaded soils. However, the specific taxa in invaded soils responsible for community differences were distinct from those in noninvaded soils that contributed to community differences. Our results indicate that, despite the phylogenetic relatedness of native and non‐native P. australis haplotypes, pathogen communities associated with the dominant non‐native haplotype are distinct from those of the rare native haplotype. Pathogen taxa that dominate either noninvaded or invaded soils suggest different potential mechanisms of invasion facilitation. These findings are consistent with the hypothesis that non‐native plant species that dominate landscapes may “cultivate” a different soil pathogen community to their rhizosphere than those of rarer native species.     

Plant invasion alters nitrogen cycling by modifying the soil nitrifying community

Plant invasions have dramatic aboveground effects on plant community composition, but their belowground effects remain largely uncharacterized. Soil microorganisms directly interact with plants and mediate many nutrient transformations in soil. We hypothesized that belowground changes to the soil microbial community provide a mechanistic link between exotic plant invasion and changes to ecosystem nutrient cycling. To examine this possible link, monocultures and mixtures of exotic and native species were maintained for 4 years in a California grassland. Gross rates of nitrogen (N) mineralization and nitrification were quantified with ¹⁵N pool dilution and soil microbial communities were characterized with DNA‐based methods. Exotic grasses doubled gross nitrification rates, in part by increasing the abundance and changing the composition of ammonia‐oxidizing bacteria in soil. These changes may translate into altered ecosystem N budgets after invasion. Altered soil microbial communities and their resulting effects on ecosystem processes may be an invisible legacy of exotic plant invasions.     

Soil physical properties associated with the invasive spotted knapweed and native grasses are similar

Centaurea maculosa Lam. (spotted knapweed), a Eurasian perennial forb, has invaded disturbed and undisturbed semiarid grasslands in the western United States. In the past, success in controlling C. maculosa and restoring invaded areas has been limited. Most research has addressed chemical aspects of invasive species interactions with soils, while potential impacts of altered soil physical properties on C. maculosa's success has not been studied. We hypothesized that the persistence of C. maculosa in semiarid rangelands might reflect an ability to alter site conditions. The objective of this study was to compare selected soil physical properties under C. maculosa-dominated and native perennial grass-dominated areas on semiarid grassland. We used six field sites in western Montana containing adjacent plots dominated by C. maculosa and by native perennial grasses. Soil physical properties including particle size fractions, bulk density, and hydraulic and thermal properties, as well as total organic carbon content, of near-surface soils were measured for each vegetation type. Soil physical properties seldom differed between C. maculosa- and native grass-dominated areas. When soil physical properties differed, the differences were inconsistent within and among sites. Presence of C. maculosa did not alter surface soil characteristics at our six sites, thus its persistence on these semi-arid grasslands cannot be explained by an ability to alter near-surface soil characteristics.     

Original Articles: Differential Influence of Plant Species on Soil Nitrogen Transformations Within Moist Meadow Alpine Tundra

Plant species can influence nitrogen (N) cycling indirectly through the feedbacks of litter quality and quantity on soil N transformation rates. The goal of this research was to focus on small-scale (within-community) variation in soil N cycling associated with two community dominants of the moist meadow alpine tundra. Within this community, the small-scale patchiness of the two most abundant species (Acomastylis rossii and Deschampsia caespitosa) provides natural variation in species cover within a relatively similar microclimate, thus enabling estimation of the effects of plant species on soil N transformation rates. Monthly rates of soil N transformations were dependent on small-scale variation in both soil microclimate and species cover. The relative importance of species cover compared with soil microclimate increased for months 2 and 3 of the 3-month growing season. Growing-season net N mineralization rates were over ten times greater and nitrification rates were four times greater in Deschampsia patches than in Acomastylis patches. Variability in litter quality [carbon:nitrogen (C:N) and phenolic:N], litter quantity (aboveground and fine-root production), and soil quality (C:N) was associated with three principal components. Variability between the species in litter quality and fine-root production explained 31% of the variation in net N mineralization rates and 36% of net nitrification rates. Site variability across the landscape in aboveground production and soil C:N explained 33% of the variation in net N mineralization rates and 21% of net nitrification rates. Within the moist meadow community, the high spatial variability in soil N transformation rates was associated with differences in the dominant species' litter quality and fine-root production. Deschampsia-dominated patches consistently had greater soil N transformation rates than did Acomastylis-dominated patches across the landscape, despite site variability in soil moisture, soil C:N, and aboveground production. Plant species appear to be an important control of soil N transformation in the alpine tundra, and consequently may influence plant community structure and ecosystem function.     

Soil Biological and Chemical Properties in Restored Perennial Grassland in California

Restoration of California native perennial grassland is often initiated with cultivation to reduce the density and cover of non‐native annual grasses before seeding with native perennials. Tillage is known to adversely impact agriculturally cultivated land; thus changes in soil biological functions, as indicated by carbon (C) turnover and C retention, may also be negatively affected by these restoration techniques. We investigated a restored perennial grassland in the fourth year after planting Nassella pulchra, Elymus glaucus, and Hordeum brachyantherum ssp. californicum for total soil C and nitrogen (N), microbial biomass C, microbial respiration, CO₂ concentrations in the soil atmosphere, surface efflux of CO₂, and root distribution (0‐ to 15‐, 15‐ to 30‐, 30‐ to 60‐, and 60‐ to 80‐cm depths). A comparison was made between untreated annual grassland and plots without plant cover still maintained by tillage and herbicide. In the uppermost layer (0‐ to 15‐cm depth), total C, microbial biomass C, and respiration were lower in the tilled, bare soil than in the grassland soils, as was CO₂ efflux from the soil surface. Root length near perennial bunchgrasses was lower at the surface and greater at lower depths than in the annual grass–dominated areas; a similar but less pronounced trend was observed for root biomass. Few differences in soil biological or chemical properties occurred below 15‐cm depth, except that at lower depths, the CO₂ concentration in the soil atmosphere was lower in the plots without vegetation, possibly from reduced production of CO₂ due to the lack of root respiration. Similar microbiological properties in soil layers below 15‐cm depth suggest that deeper microbiota rely on more recalcitrant C sources and are less affected by plant removal than in the surface layer, even after 6 years. Without primary production, restoration procedures with extended periods of tillage and herbicide applications led to net losses of C during the plant‐free periods. However, at 4 years after planting native grasses, soil microbial biomass and activity were nearly the same as the former conditions represented by annual grassland, suggesting high resilience to the temporary disturbance caused by tillage.     

Soil C and N availability determine the priming effect: microbial N mining and stoichiometric decomposition theories

The increasing input of anthropogenically derived nitrogen (N) to ecosystems raises a crucial question: how does available N modify the decomposer community and thus affects the mineralization of soil organic matter (SOM). Moreover, N input modifies the priming effect (PE), that is, the effect of fresh organics on the microbial decomposition of SOM. We studied the interactive effects of C and N on SOM mineralization (by natural ¹³C labelling adding C₄‐sucrose or C₄‐maize straw to C₃‐soil) in relation to microbial growth kinetics and to the activities of five hydrolytic enzymes. This encompasses the groups of parameters governing two mechanisms of priming effects – microbial N mining and stoichiometric decomposition theories. In sole C treatments, positive PE was accompanied by a decrease in specific microbial growth rates, confirming a greater contribution of K‐strategists to the decomposition of native SOM. Sucrose addition with N significantly accelerated mineralization of native SOM, whereas mineral N added with plant residues accelerated decomposition of plant residues. This supports the microbial mining theory in terms of N limitation. Sucrose addition with N was accompanied by accelerated microbial growth, increased activities of β‐glucosidase and cellobiohydrolase, and decreased activities of xylanase and leucine amino peptidase. This indicated an increased contribution of r‐strategists to the PE and to decomposition of cellulose but the decreased hemicellulolytic and proteolytic activities. Thus, the acceleration of the C cycle was primed by exogenous organic C and was controlled by N. This confirms the stoichiometric decomposition theory. Both K‐ and r‐strategists were beneficial for priming effects, with an increasing contribution of K‐selected species under N limitation. Thus, the priming phenomenon described in ‘microbial N mining’ theory can be ascribed to K‐strategists. In contrast, ‘stoichiometric decomposition’ theory, that is, accelerated OM mineralization due to balanced microbial growth, is explained by domination of r‐strategists.     

Local and regional assessments of the impacts of plant invaders on vegetation structure and soil properties of Mediterranean islands

Aims Although biological invasions occur throughout the world, and some invaders are widespread in many habitats, few studies on the ecological impact of invaders have examined multiple sites. We tested how the impact of three widespread plant invaders changed depending on the identity of the species and the invaded island. We also tested whether relative species loss was lower in species‐rich communities than in species‐poor ones. Location We conducted floristic surveys and soil analyses in eight Mediterranean Basin islands: Crete and Lesbos (Greece), Sardinia (Italy), Corsica, Bagaud and Porquerolles (France), and Mallorca and Menorca (Spain). Methods We compared native species richness and diversity, proportion of life forms, soil percentage nitrogen, percentage organic carbon, C/N, and soil pH in nearby paired plots of 2 × 2 m: one control and one invaded by either the deciduous tree Ailanthus altissima, the succulent subshrubs Carpobrotus spp. or the annual geophyte Oxalis pes‐caprae, across eight Mediterranean Basin islands. Results On average, the presence of invaders reduced species diversity, Carpobrotus spp. exhibiting the largest impact and Oxalis the least. However, the relative impact was island‐dependent, and was positively but weakly associated with the species richness of the recipient community. Therophytes were the life form that experienced the largest decrease across islands. The effects of invasion on soil properties were very variable. Total N changed (increased) only in plots invaded by Ailanthus, significantly decreasing the C/N ratio. The presence of this tree increased soil pH, whereas the opposite was found in plots invaded by the other two species. Organic C increased in plots invaded by Ailanthus and Carpobrotus species. Main conclusions By conducting an analysis at multiple sites, we found that the three plant invaders had an impact on plant community structure not entirely concordant with changes in soil properties. The impacts depended on the identity of the species and of the invaded island, suggesting that impact of invaders is context‐specific. The impact in terms of species loss was not lower in species‐rich than in species‐poor communities.     

Effects of exotic invasive trees on nitrogen cycling: a case study in Central Spain

We assess the hypothesis that rates of nitrogen transformations in the soil are altered upon replacement of native by exotic trees, differing in litter properties. Ailanthus altissima and Robinia pseudoacacia, two common exotic trees naturalized in the Iberian Peninsula, were compared with the native trees Ulmus minor and Fraxinus angustifolia, respectively. Naturally senesced leaves of each species were collected and C:N ratio, N and lignin content assessed. We prepared 64 litter bags per species and left them to decompose, below the canopy of the same species and below the canopy of the paired species. Dry mass, N concentration and N pool of the remaining litter were assessed after 5 and 7 months. Soil samples were collected three times during the experiment to assess soil moisture, organic matter, pH, potential mineralization rates and mineral N pools. Mineral N availability was assessed three times in the field by using ion-exchange resin-impregnated membranes. Ailanthus litter decomposed faster than Ulmus litter, probably due to the higher toughness of the latter. In spite of its high N content, Robinia litter decomposed slower than Fraxinus one, probably due to its high lignin content. In both cases, litter decomposition was faster below the exotic than the native canopies. The release of N per unit of initial litter mass was higher under both invaded situations (Ailanthus below Ailanthus and Robinia below Robinia) than under the native ones. However, soils collected below native and exotic trees neither differed in potential N mineralization rate nor in mineral N. This may be attributed to a quick plant uptake of released N and/or to a high organic matter accumulation in the soil previous to invasion that can exert a tighter control on soil N transformations than the current exotic litter.