Plant Diversity, Production, Stability, and Susceptibility to Invasion in Restored Northern Tall Grass Prairies (United States)

This study investigated the relationship among plant diversity, production, stability, and susceptibility to invasion in restored northern tall grass prairies (United States). The experiment consisted of 50 species mixtures fertilized with N or P, at high or low levels. Results from the past 5 years were as follows: (1) aboveground biomass increased and year‐to‐year variability declined with increases in plant species and functional form richness, mostly as a result of substantial increases in minimum biomass (maximum biomass was unaffected). (2) Aboveground biomass and biomass stability increased when the species in the mixture had: (a) high relative growth rates, root density, root surface area per unit of root biomass, uptake rates of N or P per unit of root surface area, and N use efficiency and (b) low root to shoot ratio. (3) Invasion of nonseeded species declined with increases in plant species and functional form richness. (4) The results from this experiment did not provide a single specific criterion for selecting an optimal species mixture. However, if the objectives of the restoration were simply to achieve an aboveground biomass variability that is less than that of growing‐season precipitation, then the seed mixtures need to have a minimum of nine species and three functional forms.     

Fertilization with nitrogen and phosphorus increases abundance of non-native species in Hawaiian montane forests

Weexamined the effects of fertilization on the diversity, abundance, and cover ofthe understory plant community of two montane wet forests in Hawaii. One siteoccupies a young substrate, where aboveground tree growth is limited bynitrogen(N), while the other site is on an older substrate, where aboveground treegrowth is limited by phosphorus (P). Both sites contained an on-going,long-termfactorial fertilization experiment in which plots were fertilized semi-annuallywith N, P, or N and P in combination. In each fertilization treatment, wemeasured density of species 0.5 m tall and percent cover ofspecies <0.5 m tall. Fertilization with N reducedspeciesrichness at the young, N-limited site, but none of the nutrient additionsaltered species richness at the older, P-limited site. Species diversity andevenness were not affected by fertilization at either site. At the site withlowN availability, plots fertilized with NP had higher densities of the non-nativeginger Hedychium gardnerianum, and at the site with lowP-availability, densities of the exotic shrub Rubusargutuswere higher in P- and NP-fertilized plots. Other effects included declines inmoss cover with fertilization at both sites, and reduced abundance of nativeseedlings in response to N and NP addition at the N-limited site. Continuedlong-term fertilization could lead to greater dominance of non-native speciesbyencouraging their growth at the expense of native species, which may sufferdecreased recruitment as fertilization and increased abundance of thenon-nativespecies may reduce suitable substrates for seedling establishment.     

Above- and belowground responses to nitrogen addition in a Chihuahuan Desert grassland

Increased available soil nitrogen can increase biomass, lower species richness, alter soil chemistry and modify community structure in herbaceous ecosystems worldwide. Although increased nitrogen availability typically increases aboveground production and decreases species richness in mesic systems, the impacts of nitrogen additions on semiarid ecosystems remain unclear. To determine how a semiarid grassland responds to increased nitrogen availability, we examined plant community structure and above- and belowground net primary production in response to long-term nitrogen addition in a desert grassland in central New Mexico, USA. Plots were fertilized annually (10 g N m⁻²) since 1995 and NPP measured from 2004 to 2009. Differences in aboveground NPP between fertilized and control treatments occurred in 2004 following a prescribed fire and in 2006 when precipitation was double the long-term average during the summer monsoon. Presumably, nitrogen only became limiting once drought stress was alleviated. Belowground NPP was also related to precipitation, and greatest root growth occurred the year following the wettest summer, decreasing gradually thereafter. Belowground production was unrelated to aboveground production within years and unrelated to nitrogen enrichment. Species richness changed between years in response to seasonal precipitation variability, but was not altered by nitrogen addition. Community structure did respond to nitrogen fertilization primarily through increased abundance of two dominant perennial grasses. These results were contrary to most nitrogen addition studies that find increased biomass and decreased species richness with nitrogen fertilization. Therefore, factors other than nitrogen deposition, such as fire or drought, may play a stronger role in shaping semiarid grassland communities than soil fertility.     

The effects of fertilization on the trait-abundance relationships in a Tibetan alpine meadow community

Aims Comparisons of the trait–abundance relationships from various habitat types are critical for community ecology, which can offer us insights about the mechanisms underlying the local community assembly, such as the relative role of neutral vs. niche processes in shaping community structure. Here, we explored the responses of trait–abundance relationships to nitrogen (N), phosphorus (P) and potassium (K) fertilization in an alpine meadow.Methods Five fertilization treatments (an unfertilized control and additions of N, P, K and NPK respectively) were implemented using randomized block design in an alpine Tibetan meadow. Species relative abundance (SRA), plant above-ground biomass and species richness were measured in each plot. For 24 common species, we measured species functional traits: saturated height, specific leaf area (SLA) and leaf dry matter content (LDMC) in each treatment but seed size only in the unfertilized control. Standard major axis (SMA) regression and phylogenetically independent contrasts (PICs) analysis were used to analyse species trait–abundance relationships in response to different fertilization treatments.Important findings Positive correlations between SRA and saturated height were raised following N, P and NPK fertilizations, which indicated an increase in light competition in these plots. In P fertilized plots, SRA was also positively correlated with LDMC because tall grasses with a nutrients conservation strategy often have a relative competitive advantage in capturing limited light and soil nutrients. In K fertilized plots, neither the trait–abundance relationships nor above-ground biomass or species richness significantly differed from that in the control, which suggests that K was not a limiting resource in our study site. These significant correlations between species traits and relative abundance in fertilized treatment suggest that trait-based selection plays an important role in determining species abundance within local communities in alpine meadows.     

Plant community and tissue chemistry responses to fertilizer and litter nutrient manipulations in a temperate grassland

Human-mediated nutrient amendments have widespread effects on plant communities. One of the major consequences has been the loss of species diversity under increased nutrient inputs. The loss of species can be functional group dependent with certain functional groups being more prone to decline than others. We present results from the sixth year of a long-term fertilization and litter manipulation study in an old-field grassland. We measured plant tissue chemistry (C:N ratio) to understand the role of plant physiological responses in the increase or decline of functional groups under nutrient manipulations. Fertilized plots had significantly more total aboveground biomass and live biomass than unfertilized plots, which was largely due to greater productivity by exotic C₃ grasses. We found that both fertilization and litter treatments affected plant species richness. Species richness was lower on plots that were fertilized or had litter intact; species losses were primarily from forbs and non-Poaceae graminoids. C₃ grasses and forbs had lower C:N ratios under fertilization with forbs having marginally greater %N responses to fertilization than grasses. Tissue chemistry in the C₃ grasses also varied depending on tissue type with reproductive tillers having higher C:N ratios than vegetative tillers. Although forbs had greater tissue chemistry responses to fertilization, they did not have a similar positive response in productivity and the number of forb species is decreasing on our experimental plots. Overall, differential nutrient uptake and use among functional groups influenced biomass production and species interactions, favoring exotic C₃ grasses and leading to their dominance. These data suggest functional groups may differ in their responses to anthropogenic nutrient amendments, ultimately influencing plant community composition.     

Density may alter diversity–productivity relationships in experimental plant communities

Contemporary biodiversity experiments, in which plant species richness is manipulated and aboveground productivity of the system measured, generally demonstrate that lowering plant species richness reduces productivity. However, we propose that community density may in part compensate for this reduction of productivity at low diversity. We conducted a factorial experiment in which plant functional group richness was held constant at three, while plant species richness increased from three to six to 12 species and community density from 440 to 1050 to 2525 seedlings m⁻². Response variables included density, evenness and above- and belowground biomass at harvest. The density gradient converged slightly during the course of the experiment due to about 10% mortality at the highest sowing density. Evenness measured in terms of aboveground biomass at harvest significantly declined with density, but the effect was weak. Overall, aboveground, belowground and total biomass increased significantly with species richness and community density. However, a significant interaction between species richness and community density occurred for both total and aboveground biomass, indicating that the diversity–productivity relationship was flatter at higher than at lower density. Thus, high species richness enabled low-density communities to reach productivity levels otherwise seen only at high density. The relative contributions of the three functional groups C₃, C₄ and nitrogen-fixers to aboveground biomass were less influenced by community density at high than at low species richness. We interpret the interaction effects between community density and species richness on community biomass by expanding findings about constant yield and size variation from monocultures to plant mixtures.     

Impact of management on biodiversity-biomass relations in Estonian flooded meadows

A negative species richness–productivity relationship is often described in grasslands at smaller scales. We aimed to study the effect of management on this relationship. In particular, we addressed the relative importance of biomass cutting, hay removal and nutrient impoverishment on species richness and growth form structure. We conducted fieldwork in flooded meadows in Alam-Pedja Nature Reserve, central Estonia. We sampled vegetation in managed and abandoned stands of two types of alluvial meadows, sedge and tall forb meadow. Aboveground biomass and litter were harvested, weighed and analysed for major plant nutrients by near infrared reflectance spectroscopy. Three groups of general additive models were developed and compared, addressing the impact of (i) productivity, (ii) nutrients and (iii) management regime on species richness. Management—mowing and hay removal—reduced the amount of litter but not aboveground biomass. Management led to a decrease in nitrogen in the biomass and enhanced species richness, particularly in the tall forb meadow. The strongest determinant of species richness was the amount of litter, exhibiting a hump-shaped relationship with richness. The effect of nitrogen supply was significant, but explained less variation. Management increased the proportion of sedges in the sedge meadow and of small herbs in the tall forb meadow. We conclude that litter removal is the most important management means to support biodiversity. On highly productive sites, reducing nutrients via hay removal is of secondary importance within a timeframe of 10 years.     

Root-shoot competition interactions cause diversity loss after fertilization: a field experiment in an alpine meadow on the Tibetan Plateau

Aims A decrease in species diversity after fertilization is a common phenomenon in grasslands; however, the mechanism causing it remains highly controversial. The light competition hypothesis to explain loss of diversity has received much attention. The aim of the present paper was to test this hypothesis.Methods Fertilization was used to control above- and belowground resources simultaneously, while shade was used to control aboveground resource in an alpine meadow on the Tibetan Plateau. Univariate general linear models was used to estimate the effects of fertilization and shade on above- and belowground vegetation characteristics, including photosynthetically active radiation (PAR) in the understory, aboveground biomass, belowground biomass, R:S ratio, species richness and Simpson's diversity index.Important findings PAR was similar in the understory of shaded and fertilized plots, but only fertilization reduced species richness and diversity, suggesting that light competition alone could not explain diversity loss after fertilization. The root biomass and R:S ratio had a significant increase in shaded plots, but the richness and diversity did not change, suggesting that root competition alone also could not explain diversity loss after fertilization in this community. Our results illustrated that the root–shoot competition interactions, investigated from a functional groups perspective, should be the most reasonable explanation leading to the diversity loss due to fertilization.     

Effects of additional illumination and fertilization on seasonal changes in fine-scale grassland community structure

Abstract. Fine-scale structure of a species-rich grassland was examined for seasonal changes caused by manipulated changes in the availability of above and below-ground resources (additional illumination with the help of mirrors and fertilization) in a field experiment. If changes induced by fertilization — which are expected to lead to a reduction in small-scale diversity —are due to intensified light competition, they should be compensated for by additional light input. Permanent plots of 40 cm × 40 cm were sampled by the point quadrat method at three angles (60°, 90° and 120° from the horizontal North-South direction), using a laser beam to position the quadrats, in early July and early September. The applied treatments did not cause apparent changes in plant leaf orientation. The degree of spatial aggregation of biomass increased seasonally in fertilized, non-illuminated plots: greater productivity at a constant light supply led to a faster growth rate of potentially dominant species, as compared to the subordinate ones. Additional illumination mitigated this effect of fertilization, indicating that the observed changes in biomass aggregation were due to increased light competition. There was a considerable seasonal decrease of variance ratio (ratio of observed variance of richness at a point and variance expected at random) in fertilized only and in illuminated only plots. In fertilized plots this was due to the positive relationship between biomass aggregation and expected variance of richness. Biomass constancy occurs to be inversely related to deficit in variance of richness. In illuminated plots, in contrast, only the observed variance of richness decreased seasonally, indicating a more uniform use of space by different species. Evidently, a deficit in variance of richness can be caused by drastically different processes, showing that the variance ratio statistic may not have a significant explanatory value in fine-scale community studies.     

Plant species richness belowground: higher richness and new patterns revealed by next-generation sequencing

Variation in plant species richness has been described using only aboveground vegetation. The species richness of roots and rhizomes has never been compared with aboveground richness in natural plant communities. We made direct comparisons of grassland plant richness in identical volumes (0.1 × 0.1 × 0.1 m) above and below the soil surface, using conventional species identification to measure aboveground richness and 454 sequencing of the chloroplast trnL(UAA) intron to measure belowground richness. We described above- and belowground richness at multiple spatial scales (from a neighbourhood scale of centimetres to a community scale of hundreds of metres), and related variation in richness to soil fertility. Tests using reference material indicated that 454 sequencing captured patterns of species composition and abundance with acceptable accuracy. At neighbourhood scales, belowground richness was up to two times greater than aboveground richness. The relationship between above- and belowground richness was significantly different from linear: beyond a certain level of belowground richness, aboveground richness did not increase further. Belowground richness also exceeded that of aboveground at the community scale, indicating that some species are temporarily dormant and absent aboveground. Similar to other grassland studies, aboveground richness declined with increasing soil fertility; in contrast, the number of species found only belowground increased significantly with fertility. These results indicate that conventional aboveground studies of plant richness may overlook many coexisting species, and that belowground richness becomes relatively more important in conditions where aboveground richness decreases. Measuring plant belowground richness can considerably alter perceptions of biodiversity and its responses to natural and anthropogenic factors.     

Consistent Effects of Biodiversity on Ecosystem Functioning Under Varying Density and Evenness

Biodiversity experiments typically vary only species richness and composition, yet the generality of their results relies on consistent effects of these factors even under varying starting conditions of density and evenness. We tested this assumption in a factorial species richness x density x evenness experiment using a pool of 60 common grassland species divided into four functional groups (grasses, legumes, tall herbs and short herbs). Richness varied from 1, 2, 4, 8 to 16 species, total planting density was 1,000 or 2,000 seeds/m², and species were sown in even or uneven proportions, where one functional group was made dominant. Aboveground plant biomass increased linearly with the logarithm of species richness in all density and evenness treatments during all three years of the experiment. This was due to a convergence of realized density and evenness within species richness levels, although functional groups which were initially made dominant retained their dominance. Between species richness levels, realized density increased, and realized evenness decreased with species richness. Thus, more individuals could coexist if they belonged to different species. Within species richness levels, higher biomass values were correlated with lower density, suggesting an underlying thinning process. However, communities with low realized evenness also had low biomass values; thus high biomass did not result from species dominance. So-called complementarity and selection effects were similar across density and evenness treatments, indicating that the mechanisms underpinning the biodiversity effects were not altered. Species richness was the dominant driver of aboveground biomass, irrespective of variations in total densities and species abundance distributions at the start of the experiment; rejecting the hypothesis that initial differences in species abundance distributions might lead to different “stable states” in community structure or biomass. Thus, results from previous biodiversity experiments that only manipulated species richness and composition should be quite robust and broadly generalizable.     

Long‐term study of root biomass in a biodiversity experiment reveals shifts in diversity effects over time

Biodiversity experiments generally report a positive effect of plant biodiversity on aboveground biomass (overyielding), which typically increases with time. Various studies also found overyielding for belowground plant biomass, but this has never been measured over time. Also, potential underlying mechanisms have remained unclear. Differentiation in rooting patterns among plant species and plant functional groups has been proposed as a main driver of the observed biodiversity effect on belowground biomass, leading to more efficient belowground resource use with increasing diversity, but so far there is little evidence to support this. We analyzed standing root biomass and its distribution over the soil profile, along a 1–16 species richness gradient over eight years in the Jena Experiment in Germany, and compared belowground to aboveground overyielding. In our long‐term dataset, total root biomass increased with increasing species richness but this effect was only apparent after four years. The increasingly positive relationship between species richness and root biomass, explaining 12% of overall variation and up to 28% in the last year of our study, was mainly due to decreasing root biomass at low diversity over time. Functional group composition strongly affected total standing root biomass, explaining 44% of variation, with grasses and legumes having strong overall positive and negative effects, respectively. Functional group richness or interactions between functional group presences did not strongly contribute to overyielding. We found no support for the hypothesis that vertical root differentiation increases with species richness, with functional group richness or composition. Other explanations, such as stronger negative plant–soil feedbacks in low‐diverse plant communities on standing root biomass and vertical distribution should be considered.     

Plant community diversity and composition affect individual plant performance

Effects of plant community diversity on ecosystem processes have recently received major attention. In contrast, effects of species richness and functional richness on individual plant performance, and their magnitude relative to effects of community composition, have been largely neglected. Therefore, we examined height, aboveground biomass, and inflorescence production of individual plants of all species present in 82 large plots of the Jena Experiment, a large grassland biodiversity experiment in Germany. These plots differed in species richness (1–60), functional richness (1–4), and community composition. On average, in more species-rich communities, plant individuals grew taller, but weighed less, were less likely to flower, and had fewer inflorescences. In plots containing legumes, non-legumes were higher and weighed more than in plots without legumes. In plots containing grasses, non-grasses were less likely to flower than in plots without grasses. This indicates that legumes positively and grasses negatively affected the performance of other species. Species richness and functional richness effects differed systematically between functional groups. The magnitude of the increase in plant height with increasing species richness was greatest in grasses and was progressively smaller in legumes, small herbs, and tall herbs. Individual aboveground biomass responses to increasing species richness also differed among functional groups and were positive for legumes, less pronouncedly positive for grasses, negative for small herbs, and more pronouncedly negative for tall herbs. Moreover, these effects of species richness differed strongly between species within these functional groups. We conclude that individual plant performance largely depends on the diversity of the surrounding community, and that the direction and magnitude of the effects of species richness and functional richness differs largely between species. Our study suggests that diversity of the surrounding community needs to be taken into account when interpreting drivers of the performance of individual plants.     

Diversity–productivity relationships in two ecologically realistic rarity–extinction scenarios

To develop a better understanding of how biodiversity loss and productivity are related, we need to consider ecologically realistic rarity (i.e. reduced evenness and increased dominance) and extinction (i.e. reduced richness) scenarios. Furthermore, we need to identify and better understand the factors that influence species and community yielding behaviors because the general conditions for overyielding are the same as those for coexistence. We established experimental tallgrass prairie plots in Iowa to determine how two ecologically realistic rarity–extinction scenarios influenced aboveground net primary productivity (ANPP) and disassembly. Equal‐mass seedlings of six tallgrass prairie species were transplanted into field plots to establish realistic declining species evenness (high, medium, low) and richness (4, 1) treatments. Across declining evenness treatments, the relative abundance of the ubiquitous tall species Andropogon gerardii increased, the relative abundance of the tall species Salvia azurea was constant, and the relative abundance of two short (dissimilar height scenario) or two tall species (tall scenario) decreased. Monocultures of Andropogon represented a continuation of this trend until there was complete dominance by Andropogon and extinction of all other species. Our treatments also allowed us to test if variation in plant height contributes to the complementarity effect. Niche partitioning in plant height was not positively related to complementarity. The effects of declining species evenness and richness on the diversity–productivity relationship were different for these two ecologically realistic rarity–extinction scenarios. Specifically, as diversity declined across treatments, ANPP and the selection effects decreased in tall communities, but not in dissimilar communities. Additionally, differences between these two scenarios revealed that decreased species yielding behavior is associated with two tallgrass prairie extinction risk factors, rarity and short height. The differences between these scenarios demonstrate the importance of incorporating the known patterns of diversity declines into future studies.     

Species richness loss after nutrient addition as affected by N:C ratios and phytohormone GA3 contents in an alpine meadow community

Aims From the light-competition hypothesis, competition for light is asymmetric and the observed increases in plant-size variability with increasingly denser canopies are primarily due to competition for light. Greater plant height provides pre-emptive access to light and produces increased height differences among species. The question is what produces these differences in plant height or height growth response among species in response to fertilization.Methods In 2009, a field experiment of N, P and N + P enrichments at three levels each was initiated in an alpine meadow on the northeast Qinghai-Tibet Plateau. Effects of fertilization on species richness, aboveground net primary production (ANPP), relative light intensity and plant height of different plant functional groups were determined. Festuca ovina (grass), Kobresia humilis (sedge), Oxytropis ochrocephala (legume), Taraxacum lugubre (rosette forb) and Geranium pylzowianum (upright forb) were selected as exemplars of each of the indicated functional groups. The N:C ratios in aboveground biomass, gibberellic acid (GA 3) concentrations in leaves, plant heights and height relative growth rate (RGR) of these exemplar species were analyzed in detail.Important findings Species richness of grasses significantly increased with increasing N + P levels. Species richness of legumes and upright forbs decreased after N and N + P additions. P addition had no significant effect on species richness. The effects of N + P addition on species richness and ANPP were consistently stronger than those of the single N or P fertilization. Reductions in species richness caused by nutrient addition paralleled the increases in ANPP and decreases in light intensity under the canopies, indicating indirect effect of nutrient addition on species richness via ANPP-induced light competition. The exemplar species that responded most positively to fertilization in height and RGR also displayed stronger increases in their GA 3 content and N:C ratios. GA 3 concentrations and N:C ratios were positively correlated with height RGR when the data were pooled for all species. The tallest and the fastest-growing grass, F. ovina, had the largest increase in N:C ratios and the highest leaf GA 3 concentrations after nutrient addition. These results indicated that differential responses of GA 3 concentrations and N:C ratios to fertilization were related to the inequality in plant heights among species.     

模拟降雨量变化对黄河三角洲湿地植物群落特征的影响

气候变化背景下,降雨变化能够深刻影响河口湿地土壤水盐条件,而土壤水盐条件是影响植物群落特征的关键环境因子。本研究以黄河三角洲湿地植物群落为对象,依托野外降雨控制试验平台(减雨60%、减雨40%、自然对照、增雨40%、增雨60%),探讨了经过6年降雨处理后湿地植物群落特征对降雨量变化的响应及机制。结果表明: 随降雨量增加,土壤电导率显著降低,土壤湿度显著增大。降雨量变化影响了植物群落物种组成,增雨处理降低了碱蓬和盐地碱蓬的优势地位,提高了荻和白茅的优势地位。随降雨量增加,植物群落Shannon指数和Margalef丰富度指数显著提高。与对照相比,增减雨处理均降低了群落频度、多度和盖度,增雨60%处理群落频度显著降低54.9%,减雨60%、减雨40%、增雨40%、增雨60%处理群落多度分别显著降低38.9%、33.8%、35.8%和45.7%。随降雨量增加,植物群落地上生物量显著增加,但可能受淹水胁迫的影响,增雨60%处理地上生物量显著低于增雨40%。Margalef丰富度指数与地上生物量呈显著正相关;地上生物量、Shannon指数、Margalef丰富度指数、Simpson多样性指数均与土壤电导率呈显著负相关;地上生物量与土壤湿度呈显著正相关。降雨量变化通过改变黄河三角洲湿地土壤水盐条件显著影响了植物群落生长特征、物种组成和多样性。     

光照，施肥及刈割对垂穗披碱草生长的影响

以甘南高寒草甸常见牧草垂穗披碱草（Elymus nutans）为研究对象,比较不同光照、肥力条件下,垂穗披碱草对刈割的生长和补偿反应。研究发现,光照和肥力存在交互作用。施肥、中度光照下,未刈割单株垂穗披碱草地上生物量最大,为3．239g。施肥、中度光照下,比较刈割植株地上生物量补偿发现：晚期刈割单株地上生物量的补偿大于早期刈割;刈割强度无显著影响;刈割两次单株地上生物量的补偿最大。施肥、中度光照、晚期、轻度、一次刈割时,单株垂穗披碱草地上生物量累积最大,为4．500g。全光照、未施肥条件下,刈割对单株地上生物量的补偿无影响。就补偿机制而言,植株地上生物量的增加以地下生物量为代价。     

The unseen invaders: introduced earthworms as drivers of change in plant communities in North American forests (a meta‐analysis)

Globally, biological invasions can have strong impacts on biodiversity as well as ecosystem functioning. While less conspicuous than introduced aboveground organisms, introduced belowground organisms may have similarly strong effects. Here, we synthesize for the first time the impacts of introduced earthworms on plant diversity and community composition in North American forests. We conducted a meta‐analysis using a total of 645 observations to quantify mean effect sizes of associations between introduced earthworm communities and plant diversity, cover of plant functional groups, and cover of native and non‐native plants. We found that plant diversity significantly declined with increasing richness of introduced earthworm ecological groups. While plant species richness or evenness did not change with earthworm invasion, our results indicate clear changes in plant community composition: cover of graminoids and non‐native plant species significantly increased, and cover of native plant species (of all functional groups) tended to decrease, with increasing earthworm biomass. Overall, these findings support the hypothesis that introduced earthworms facilitate particular plant species adapted to the abiotic conditions of earthworm‐invaded forests. Further, our study provides evidence that introduced earthworms are associated with declines in plant diversity in North American forests. Changing plant functional composition in these forests may have long‐lasting effects on ecosystem functioning.     

Climate and species affect fine root production with long-term fertilization in acidic tussock tundra near Toolik Lake,Alaska

Long-term fertilization of acidic tussock tundra has led to changes in plant species composition, increases in aboveground production and biomass and substantial losses of soil organic carbon (SOC). Root litter is an important input to SOC pools, although little is known about fine root demography in tussock tundra. In this study, we examined the response of fine root production and live standing fine root biomass to short- and long-term fertilization, as changes in fine root demography may contribute to observed declines in SOC. Live standing fine root biomass increased with long-term fertilization, while fine root production declined, reflecting replacement of the annual fine root system of Eriophorum vaginatum, with the long-lived fine roots of Betula nana. Fine root production increased in fertilized plots during an unusually warm growing season, but remained unchanged in control plots, consistent with observations that B. nana shows a positive response to climate warming. Calculations based on a few simple assumptions suggest changes in fine root demography with long-term fertilization and species replacement could account for between 20 and 39% of the observed declines in SOC stocks.     

Response of Alpine Plants to Nitrogen Addition on the Tibetan Plateau: A Meta-analysis

To identify the general effects of nitrogen addition on alpine plants, we used a meta-analysis approach to synthesize 599 observations from 51 studies on the Tibetan Plateau. Nitrogen addition significantly increased plant height by 19.0 %, plant biomass by 29.7 %, graminoid aboveground biomass by 89.8 %, and sedge aboveground biomass by 75.6 % but significantly decreased legume aboveground biomass by 34.5 %, forb aboveground biomass by 23.8 %, and species richness by 11.2 %. The effect of nitrogen addition on aboveground plant biomass and plant height increased with increasing the nitrogen addition rate. The effect of nitrogen addition on plant height increased with increasing mean annual precipitation but decreased with increasing mean annual temperature. Our findings suggested that the effect of nitrogen addition on alpine plants varied with plant functional types and nitrogen addition rate. In addition, climatic warming and precipitation changes may regulate the response of alpine plants to nitrogen addition on the Tibetan Plateau.