Negative plant–soil feedback and species coexistence

Different mechanisms, including equilibrium and non-equilibrium processes, have been taken into account as possible theoretical explanations of species coexistence. Despite the ample evidence on the existence of negative plant–soil feedback in both agriculture and natural vegetation, the role of these processes in the organization and dynamics of plant communities has so far been neglected. In this study, simulations by an individual-based competition model show how the intensity of negative feedback on individual plant performance can produce faster successional dynamics and allow species coexistence in two- and multi-species systems. The results show that even low levels of negative plant–soil feedback can enable species coexistence and often produce cyclic population dynamics. Moreover, the model highlights how negative feedback can generate positive reciprocal interspecific interactions at the population level, despite the fact that only competitive interactions is present between individual plants. In fact, competitive effects occur on a short-term scale, but positive reciprocal species interactions emerge only if negative feedback affects all species and if longer periods of simulation, more than the species life span, are considered. An important outcome of the model is the evidence that the effects at population level are timescale-dependent, thus showing the limitation of short-term species removal experiments used in traditional competition studies.     

Bird communities in habitats along a successional gradient: Divergent patterns of species richness,specialization and threat

The long history of human influence on northern temperate landscapes has created a mosaic of successional stages, from closed forest to open grassland. Various species thus adapted to different habitats and it is interesting to explore how these differences in species composition among particular successional stages translate into differences at the community level. For this purpose, we surveyed breeding birds in 233 patches of five different habitats covering a gradient from bare ground to forest in 29 abandoned military training sites scattered throughout the Czech Republic. Linear mixed effects modelling revealed that late-successional habitats (dense scrubland and forest) were the most species-rich, whereas early-successional stages hosted bird communities with the highest habitat specialization and threat level. These results suggest that the habitats of late-successional stages are important for the maintenance of high bird species richness, but that early-successional habitats are essential for highly specialized and threatened bird species. Given the highly adverse impacts of agricultural intensification and land abandonment on open habitats, it is necessary to promote factors creating initial successional stages suitable for specialized and threatened species.     

森林次生演替和土壤层次对微生物群落结构的影响

森林次生演替与生态系统结构和功能的动态变化密切相关。大多数研究主要关注植物群落以及土壤有机碳(SOC)的变化,然而土壤微生物群落如何响应森林次生演替还需要进一步探究。本研究以长白山森林次生演替序列(20、80、120、200和≥300年)和两个土壤层次为对象,采用磷脂脂肪酸微生物标志物,探究温带森林次生演替过程中地下微生物群落结构变化。森林次生演替改变了土壤微生物群落结构,主要归因于某些特定微生物类群的变化,演替前期革兰氏阴性菌和腐生真菌占主导,而在演替后期革兰氏阳性菌和丛枝菌根真菌占主导。另外,土壤有机质数量和质量差异是影响微生物群落结构和生物量的主要环境因素。森林演替前期和中期增加的SOC含量促进了微生物生物量,而演替后期增加的难分解芳香族有机组分抑制了微生物生物量合成。土壤层次间理化性质的差异导致微生物群落变化,有机质层高的SOC以及氮含量导致更多微生物生物量的合成。微生物群落在时间和空间尺度的变化及其驱动因素反映了生态系统结构和功能对环境变化的响应。     

Phylogenetic conservatism in plant‐soil feedback and its implications for plant abundance

We examined whether plant‐soil feedback and plant‐field abundance were phylogenetically conserved. For 57 co‐occurring native and exotic plant species from an old field in Canada, we collected a data set on the effects of three soil biota treatments on plant growth: net whole‐soil feedback (combined effects of mutualists and antagonists), feedback with arbuscular mycorrhizal fungi (AMF) collected from soils of conspecific plants, and feedback with Glomus etunicatum, a dominant mycorrhizal fungus. We found phylogenetic signal in both net whole‐soil feedback and feedback with AMF of conspecifics; conservatism was especially strong among native plants but absent among exotics. The abundance of plants in the field was also conserved, a pattern underlain by shared plant responses to soil biota. We conclude that soil biota influence the abundance of close plant relatives in nature.     

Above- and belowground effects of plant-soil feedback from exotic Solidago canadensis on native Tanacetum vulgare

Plant-soil feedback responses for native and invasive plant species are well documented, but little is known about how feedback effects from the soil biota community affect plant interactions with herbivores. Here we examine whether changes of the soil biota community by the successful invader Solidago canadensis influence growth and herbivore susceptibility of two coexisting native plant species (Tanacetum vulgare, Melilotus albus). Root zone soil from two different habitat types (‘urban’ and ‘suburban’) was collected and used as inocula in a plant-soil feedback study. Each plant species was grown either in its own soil biota community or with the community with a history from the competitive invasive or native plant species. To identify potential drivers of responses to the different soil biota communities, we analyzed root colonization by arbuscular mycorrhizal fungi and dark-septate endophytes (DSE), and the community composition of soil inhabiting nematodes at the end of our experiment. Results show that S. canadensis and M. albus were not affected by soil history. In contrast, T. vulgare showed increased plant growth in ‘foreign’ soil derived from S. canadensis root zone compared with its ‘home’ soil suggesting a growth promotion by the soil biota community of S. canadensis. From the examined drivers, the abundance of DSE explained the growth response of T. vulgare to the S. canadensis soil biota community best. However, shoot herbivory by banded snails (Cepaea nemoralis, C. hortensis) was not affected by soil history, but by the habitat type where the soil inocula originated. Our study shows that a native plant species may profit from the presence of an invasive competitor mediated by changes in the soil biota community.     

Inoculation with a Native Soil Community Advances Succession in a Grassland Restoration

Restoration on post‐agricultural land may be hindered by the degradation of the soil community, which has been shown to contribute to structuring plant communities and driving succession. Our experiment tested the effect of inoculation with remnant grassland whole soil with or without nurse plants on the survival and growth of uninoculated early and late successional plant species. In 2007 and 2008, we planted uninoculated early, mid, and late successional plant species 0.25–2 m away from a central point of inoculated nurse plants. We found a negative response to inoculation on early successional plants and a positive response to inoculation on mid to late successional plants. This work suggests that the restoration of the soil community is critical to establishing a late successional plant community and that the benefit of inoculated plants can spread to neighbors.     

Arbuscular mycorrhizal fungal communities change among three stages of primary sand dune succession but do not alter plant growth

Plant interactions with soil biota could have a significant impact on plant successional trajectory by benefiting plants in a particular successional stage over others. The influence of soil mutualists such as mycorrhizal fungi is thought to be an important feedback component, yet they have shown benefits to both early and late successional plants that could either retard or accelerate succession. Here we first determine if arbuscular mycorrhizal (AM) fungi differ among three stages of primary sand dune succession and then if they alter growth of plants from particular successional stages. We isolated AM fungal inoculum from early, intermediate or late stages of a primary dune succession and compared them using cloning and sequencing. We then grew eight plant species that dominate within each of these successional stages with each AM fungal inoculum. We measured fungal growth to assess potential AM functional differences and plant growth to determine if AM fungi positively or negatively affect plants. AM fungi isolated from early succession were more phylogenetically diverse relative to intermediate and late succession while late successional fungi consistently produced more soil hyphae and arbuscules. Despite these differences, inocula from different successional stages had similar effects on the growth of all plant species. Host plant biomass was not affected by mycorrhizal inoculation relative to un‐inoculated controls. Although mycorrhizal communities differ among primary dune successional stages and formed different fungal structures, these differences did not directly affect the growth of plants from different dune successional stages in our experiment and therefore may be less likely to directly contribute to plant succession in sand dunes.     

Do forest soil microbes have the potential to resist plant invasion? A case study in Dinghushan Biosphere Reserve (South China)

Successful invaders must overcome biotic resistance, which is defined as the reduction in invasion success caused by the resident community. Soil microbes are an important source of community resistance to plant invasions, and understanding their role in this process requires urgent investigation. Therefore, three forest communities along successional stages and four exotic invasive plant species were selected to test the role of soil microbes of three forest communities in resisting the exotic invasive plant. Our results showed that soil microbes from a monsoon evergreen broadleaf forest (MEBF) (late-successional stage) had the greatest resistance to the invasive plants. Only the invasive species Ipomoea triloba was not sensitive to the three successional forest soils. Mycorrhizal fungi in early successional forest Pinus massonina forest (PMF) or mid-successional forest pine-broadleaf mixed forest (PBMF) soil promoted the growth of Mikania micrantha and Eupatorium catarium, but mycorrhizal fungi in MEBF soil had no significant effects on their growth. Pathogens plus other non-mycorrhizal microbes in MEBF soil inhibited the growth of M. micrantha and E. catarium significantly, and only inhibited root growth of E. catarium when compared with those with mycorrhizal fungi addition. The study suggest that soil mycorrhizal fungi of early-mid-successional forests benefit invasive species M. micrantha and E. catarium, while soil pathogens of late-successional forest may play an important role in resisting M. micrantha and E. catarium. The benefit and resistance of the soil microbes are dependent on invasive species and related to forest succession. The study gives a possible clue to control invasive plants by regulating soil microbes of forest community to resist plant invasion.     

Plant-soil feedback as a mechanism of invasion by Carpobrotus edulis

Invasive plant species have been suggested to change the composition of the soil community in a way that results in a positive feedback for them and a negative feedback for the native plant community. Carpobrotus edulis, a species native to South Africa, is one of the most aggressive exotic species in Mediterranean Europe. Although several aspects of its invasion biology have been studied, the occurrence of plant-soil feedback has been scarcely investigated. We first checked for the existence of biotic resistance in soils from two invaded sites of Mediterranean Europe and one site in the native area. Secondly, we evaluated the effects of soil conditioning on the germination and plant growth of C. edulis and two key species of native dunes. Finally, we tested the effects of short- and long-term soil conditioning on the performance and reproductive effort of C. edulis. Our results show that at first there is a natural resistance to invasion by the soil biota. Later, biotic resistance in invaded soil is suppressed by the establishment of a soil community that enhances the growth of C. edulis and that negatively influences the growth and survival of the native plants. Long-term soil conditioning in the field resulted in shifts in the balance between vegetative growth and sexual reproduction. Long-term invasion was also reflected in high levels of endophyte colonization by chytrids in roots, although the physiological consequences of this colonization remain unknown. The results obtained illustrate a mechanism that explains how C. edulis breaks the initial biotic resistance of newly-invaded landscapes. Finally, this study highlights the importance of studying plant-soil interactions on different members of the plant community and temporal stages in order to fully understand invasion.     

SPECIES-AREA RELATIONSHIP DURING PRIMARY SUCCESSION IN GRANITE OUTCROP PLANT COMMUNITIES

The species-area relationship and its underlying explanatory mechanisms were investigated in a primary successional sere on a southeastern (United States) granite outcrop. There, plant communities occupy soil-filled depressions separated from one another by areas of bare rock. They have been termed “island communities.” Soil depth and area increase as succession proceeds from Sedum smallii, to lichen-annual, to annual-perennial, and to herb-shrub-tree stages. Although plant species richness is significantly and positively correlated with island area in the system studied (all successional stages considered), the relationship between species richness and island maximum soil depth is stronger. However, island maximum soil depth and area are significantly and positively correlated. The exponential function describes the speciesarea relationship better than the transformed power function. Within successional stage, species richness shows no significant relationship with area or depth, except for late-successional island communities. Processes related to community successional development may explain the species-area relationship that exists when islands of all stages are considered. However, mechanisms related to equilibrium between extinction and immigration may be responsible for the speciesarea relationship for late-successional island communities.     

Plant-soil feedbacks: a meta-analytical review

Plants can change soil biology, chemistry and structure in ways that alter subsequent plant growth. This process, referred to as plant–soil feedback (PSF), has been suggested to provide mechanisms for plant diversity, succession and invasion. Here we use three meta-analytical models: a mixed model and two Bayes models, one correcting for sampling dependence and one correcting for sampling and hierarchical dependence (delta-splitting model) to test these hypotheses. All three models showed that PSFs have medium to large negative effects on plant growth, and especially grass growth, the life form for which we had the most data. This supports the hypothesis that PSFs, through negative frequency dependence, maintain plant diversity, especially in grasslands. PSFs were also large and negative for annuals and natives, but the delta-splitting model indicated that more studies are needed for these results to be conclusive. Our results support the hypotheses that PSFs encourage successional replacements and plant invasions. Most studies were performed using monocultures of grassland species in greenhouse conditions. Future research should examine PSFs in plant communities, non-grassland systems and field conditions.     

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.     

Enhancement of late successional plants on ex-arable land by soil inoculations

Restoration of species-rich grasslands on ex-arable land can help the conservation of biodiversity but faces three big challenges: absence of target plant propagules, high residual soil fertility and restoration of soil communities. Seed additions and top soil removal can solve some of these constraints, but restoring beneficial biotic soil conditions remains a challenge. Here we test the hypotheses that inoculation of soil from late secondary succession grasslands in arable receptor soil enhances performance of late successional plants, especially after top soil removal but pending on the added dose. To test this we grew mixtures of late successional plants in arable top (organic) soil or in underlying mineral soil mixed with donor soil in small or large proportions. Donor soils were collected from different grasslands that had been under restoration for 5 to 41 years, or from semi-natural grassland that has not been used intensively. Donor soil addition, especially when collected from older restoration sites, increased plant community biomass without altering its evenness. In contrast, addition of soil from semi-natural grassland promoted plant community evenness, and hence its diversity, but reduced community biomass. Effects of donor soil additions were stronger in mineral than in organic soil and larger with bigger proportions added. The variation in plant community composition was explained best by the abundances of nematodes, ergosterol concentration and soil pH. We show that in controlled conditions inoculation of soil from secondary succession grassland into ex-arable land can strongly promote target plant species, and that the role of soil biota in promoting target plant species is greatest when added after top soil removal. Together our results point out that transplantation of later secondary succession soil can promote grassland restoration on ex-arable land.     

Effects of canopy gaps on the growth of tree seedlings from subtropical broad-leaved evergreen forests of southern China

Seedling performance, morphology, and leaf characteristics of evergreen tree species from different successional stages of Chinese broadleaved evergreen forests were studied in four simulated gap environments: in 100, 55, 33, and 18% of full sunlight. The first two represent the light regimes of clearings (gaps secondarily become more open due to human activity and natural erosion). The hypothesis was tested that early-successional species achieve their greatest seedling size in clearings and contrast in this respect with late-successional species. Late-successional species were expected to show a stronger morphological response (leaf/root ratio) to clearings than early-successional ones. The results provided some evidence in support of these hypotheses with one exception, namely that late-successional Castanopsis fargesii appeared to be a highly sun-tolerant species. It is suggested that Castanopsis seedlings are competitive in large gaps and clearings.     

植物-土壤反馈与草地群落演替:菌根真菌和土壤病原菌的调控作用

由植物引起的根际土壤生物或非生物环境的改变能够反馈影响群落中不同植物的生长,直接改变共存植物的相对竞争关系,推动群落结构的动态变化。作为土壤生物群落的重要组成部分,土壤微生物在植物-土壤反馈关系中起到重要的调控作用,对解释植物群落的演替进程和方向有着重要的意义。在草地植物群落演替的早期阶段和外来物种入侵的过程中,宿主植物对丛枝菌根真菌（AMF）的依赖性较低,受本地病原菌的影响较小,一般不存在负反馈。在演替后期,植物对AMF更具依赖性,而积累的病原菌则产生较强的负反馈效应,从而促进群落物种共存和植物多样性,提高草地生产力和稳定性。研究微生物-植物反馈机制不仅有助于完善草地退化与恢复理论,还对退化草地恢复治理的实践有着指导意义。未来关于根际微生物-植物反馈在草地群落演替中的作用应该加强以下几方面的研究：（1）在实验方法上,开展专性微生物-植物反馈研究;（2）在测定指标上,进一步量化不同微生物在反馈关系中的功能差异;（3）在研究对象上,加强土壤微生物在植物群落水平的反馈研究;（4）在应用上,明晰植物-土壤反馈在退化草地恢复过程中的作用,指导草地管理实践。     

Intraspecific and intergenerational differences in plant–soil feedbacks

Interactions between plant and soil communities are known to play an integral role in shaping ecosystems. Plants influence the composition of soil communities and soil communities in turn influence plant performance. Such a plant–soil feedback may incur selection pressure on plants and the associating soil community. However, the evolutionary consequences of these above–belowground feedback interactions remain largely speculative. Here we assess whether plant–soil feedback effects differ between intraspecific plant populations and between generations within the same plant population. We used two populations of Trifolium pratense and assessed their performance when grown in association with their home versus away soil biota. Both populations were colonized by distinct microbial communities and performed better with their own home soil communities than with the soil community from the other intraspecific population, demonstrating intraspecific positive feedback effects of home soil. In one of the two populations, we found that plant performance and the root associated microbiota community differed between parental and progeny plants when inoculated with their own home soil. Differences in root associated community characteristics could explain more than 80% of the variation in performance among the progeny and parental plants. Our results highlight that intraspecific differences in both plant and associated soil communities shape plant–soil feedback effects, and consequently indicate that plant–soil feedback can influence the direction of selection between intraspecific plant populations.     

Do plant parasitic nematodes have differential effects on the productivity of a fast- and a slow-growing grass species?

We have examined the interaction between plant parasitic nematodes and plant species from different stages of grassland succession. In these grasslands, fertiliser application was stopped in order to restore the former nutrient-poor ecosystems. This management resulted in a reversed succession of high- to low-productivity. Nematodes isolated from a high-productive early-successional field and a low-productive late-successional field were inoculated to sterilised soil planted with seedlings of either Lolium perenne (a fast-growing early-successional species) or Festuca rubra (a slow-growing late-successional species). The experiment was performed at low and high supply rates of nutrients. We hypothesised that at a low nutrient supply rate the growth of L. perenne will be more reduced by nematode herbivory than the growth of F. rubra. Furthermore, we hypothesised that higher numbers of plant parasitic nematodes will develop under L. perenne. We found no support for our first hypothesis, because nematodes did not affect plant growth. Our results suggest that changes in the nutrient availability rather than plant parasitic nematodes affect plant succession in impoverished grasslands. On the other hand, plant species and nutrient supply rate significantly affected the density and composition of the plant parasitic nematode community. In line with our second hypothesis, plant parasitic nematodes reproduced better on the fast-growing L. perenne than on the slow-growing F. rubra. Our results, therefore, suggest that the succession of the plant parasitic nematode community is probably more affected by changes in the plant community than the other way round.     

Punching above their weight: low-biomass non-native plant species alter soil properties during primary succession

Non‐native invasive plants can greatly alter community and ecosystem properties, but efforts to predict which invasive species have the greatest impacts on these properties have been generally unsuccessful. An hypothesis that has considerable promise for predicting the effects of invasive non‐native plant species is the mass ratio hypothesis (i.e. that dominant species exert the strongest effects). We tested this hypothesis using data from a four year removal experiment in which the presence of two dominant shrub species (one native and the other not), and subordinate plant species, were manipulated in factorial combinations over four years in a primary successional floodplain system. We measured the effects of these manipulations on the plant community, soil nutrient status and soil biota in different trophic levels of the soil food web. Our experiment showed that after four years, low‐biomass non‐native plant species exerted disproportionate belowground effects relative to their contribution to total biomass in the plant community, most notably by increasing soil C, soil microbial biomass, altering soil microbial community structure and increasing the abundance of microbial‐feeding and predatory nematodes. Low‐biomass, non‐native plant species had distinct life history strategies and foliar traits (higher foliar N concentrations and higher leaf area per unit mass) compared with the two dominant shrub species (97% of total plant mass). Our results have several implications for understanding species’ effects in communities and on soil properties. First, high‐biomass species do not necessarily exert the largest impacts on community or soil properties. Second, low‐biomass, inconspicuous non‐native species can influence community composition and have important trophic consequences belowground through effects on soil nutrient status or resource availability to soil biota. Our finding that low‐biomass non‐native species influence belowground community structure and soil properties more profoundly than dominant species demonstrates that the mass ratio hypothesis does not accurately predict the relative effects of different coexisting species on community‐ and ecosystem‐level properties.     

High-nitrogen and low-irradiance can restrict energy utilization in photosynthesis of successional tree species in low subtropical forest

Responses of photosynthesis and the partition of energy utilization to high-nitrogen importation and high-light intensity in leaves of three dominant tree species of subtropical forest,including sun plant or early-successional species Schima superba,mesophyte or intermediate-successional species Canstanopsis hystrix,and shading-tolerant plant or late-successional species Cryptocarya concinna were studied by using the CO2 exchange system and chlorophyll fluorescence method.Our results showed that,regardless of plant species,net photosynthetic rate(Pn)was higher in high-nitrogen supply and high irradiance(HNHL)plants than in low-nitrogen supply and high irradiance(LNHL)plants,implying that low-nitrogen importation would limit Pn of plants grown under high irradiance.However, high-nitrogen supply and low irradiance(HNLL)plants had a lower Pn.Insignificant change of quantum yield(Fv′/Fm′)in opened PS II was found in leaves of HNHL,LNHL or HNLL plants of S.superba and C. hystrix,while a higher Fv′/Fm′occurred in HNHL plants of C.concinna in comparison with LNHL or HNLL plants.The HNHL plants of C.concinna also had a higher photochemical quantum yield(△F/Fm′) than LNHL or HNLL plants,however no similar responses were found in plants of S.superba and C. hystrix(P<0.05).In the irradiance range of 0―2000μmol photon·m -2·s -1,the fraction of energy consumed by photochemistry(φ PSII )was 18.2%in LNHL plants of S.superba which was higher than that in HNHL plants(P>0.05)and it was significantly higher than in HNLL plants(P<0.05).C.hystrix also had a similar response inφ PSII to nitrogen supply and irradiance.Regardless of species HNLL plants had a significantφ PSII and higher heat dissipation in light,and this effect was more severe in C.concinna than in S.superba or C.hystrix.The results may mean that high-nitrogen importation by nitrogen deposit and low irradiance caused by changing climate or air pollution would more severely restrict photosynthetic processes in the late-successional species C.concinna than in the early-successional species S.superba and intermediate-successional species C.hystrix.The continuous high-nitrogen precipitation in the future and the over cast mist or pollution smoke could induce late-successional species to degrade,however,early-successional species would be more adapted to competition for more resources to keep their dominance in ecosystems.In this sense,the zonal vegetation may accelerate degradation caused by high nitrogen precipitation and low irradiance,while the early-successional and mesophytic vegetations can remain longer.Thus,nitrogen precipitation may play an important role in plant community succession.     

Accumulation of local pathogens: a new hypothesis to explain exotic plant invasions

Recent studies have concluded that release from native soil pathogens may explain invasion of exotic plant species. However, release from soil enemies does not explain all plant invasions. The invasion of Ammophila arenaria (marram grass or European beach grass) in California provides an illustrative example for which the enemy release hypothesis has been refuted. To explore the possible role of plant–soil community interactions in this invasion, we developed a mathematical model. First, we analyzed the role of plant–soil community interactions in the succession of A. arenaria in its native range (north-western Europe). Then, we used our model to explore for California how alternative plant–soil community interactions may generate the same effect as if A. arenaria were released from soil enemies. This analysis was carried out by construction of a 'recovery plane' that discriminates between plant competition and plant–soil community interactions. Our model shows that in California, the accumulation of local pathogens by A. arenaria could result in exclusion of native plant species. Moreover, this mechanism could trigger the rate and spatial pattern of invasive spread generally observed in nature. We propose that our 'accumulation of local pathogens' hypothesis could serve as an alternative explanation for the enemy release hypothesis to be considered in further experimental studies on invasive plant species.