Janzen-Connell effects are widespread and strong enough to maintain diversity in grasslands

Crop rotation schemes are believed to work by preventing specialist soil-borne pests from depressing the future yields of similar crops. In ecology, such negative plant-soil feedbacks may be viewed as a type of Janzen-Connell effect, which promotes species coexistence and diversity by preventing the same species from repeatedly occupying a particular site. In a controlled greenhouse experiment with 24 plant species and using soils from established field monocultures, we reveal community-wide soil-based Janzen-Connell effects between the three major functional groups of plants in temperate European grasslands. The effects are much stronger and more prevalent if plants are grown in interspecific competition. Using several soil treatments (gamma irradiation, activated carbon, fungicide, fertilizer) we show that the mechanism of the negative feedback is the buildup of soil pathogens which reduce the competitive ability of nearly all species when grown on soils they have formerly occupied. We further show that the magnitude of the change in competitive outcome is sufficient to stabilize observed fitness differences between functional groups in reasonably large communities. The generality and strength of this negative feedback suggests that Janzen-Connell effects have been underestimated as drivers of plant diversity in temperate ecosystems.     

Specificity between Neotropical tree seedlings and their fungal mutualists leads to plant-soil feedback

A growing body of evidence obtained largely from temperate grassland studies suggests that feedbacks occurring between plants and their associated soil biota are important to plant community assemblage. However, few studies have examined the importance of soil organisms in driving plant-soil feedbacks in forested systems. In a tropical forest in central Panama, we examined whether interactions between tree seedlings and their associated arbuscular mycorrhizal fungi (AMF) lead to plant-soil feedback. Specifically, do tropical seedlings modify their own AMF communities in a manner that either favors or inhibits the next cohort of conspecific seedlings (i.e., positive or negative feedback, respectively)? Seedlings of two shade-tolerant tree species (Eugenia nesiotica, Virola surinamensis) and two pioneer tree species (Luehea seemannii, Apeiba aspera) were grown in pots containing identical AMF communities composed of equal amounts of inoculum of six co-occurring AMF species. The different AMF-host combinations were all exposed to two light levels. Under low light (2% PAR), only two of the six AMF species sporulated, and we found that host identity did not influence composition of AMF spore communities. However, relative abundances of three of the four AMF species that produced spores were influenced by host identity when grown under high light (20% PAR). Furthermore, spores of one of the AMF species, Glomus geosporum, were common in soils of Luehea and Eugenia but absent in soils of Apeiba and Virola. We then conducted a reciprocal experiment to test whether AMF communities previously modified by Luehea and Apeiba differentially affected the growth of conspecific and heterospecific seedlings. Luehea seedling growth did not differ between soils containing AMF communities modified by Luehea and Apeiba. However, Apeiba seedlings were significantly larger when grown with Apeiba-modified AMF communities, as compared to Apeiba seedlings grown with Luehea-modifed AMF communities. Our experiments suggest that interactions between tropical trees and their associated AMF are species-specific and that these interactions may shape both tree and AMF communities through plant-soil feedback.     

Soil biota drive expression of genetic variation and development of population-specific feedbacks in an invasive plant

Invasive plant species alter soils in ways that may affect the success of subsequent generations, creating plant-soil feedbacks. Ailanthus altissima is an invasive tree introduced two centuries ago to North America. We hypothesized that geographically distinct populations of A. altissima have established feedbacks specific to their local environment, due to soil communities cultivated by A. altissima. We collected seeds and soils from three populations in the eastern United States, and in the greenhouse reciprocally planted all families in all collected soils as well as in a control mixed soil, and in soils that had been irradiated for sterilization. There were positive plant-soil feedbacks for two populations in the live field-collected soils, but strong negative feedbacks for the third population. There were no population-level performance differences or feedbacks in the sterilized population locale soils, supporting a soil biotic basis for feedbacks and for the expression of genetic differentiation in A. altissima. If populations of Ailanthus altissima vary in the extent to which they benefit from and promote these plant-soil biota feedbacks, the interaction between invader and invaded community may be more important in determining the course of invasion than are the characteristics of either alone.     

Plant-soil feedback: testing the generality with the same grasses in serpentine and prairie soils

Plants can alter soil properties in ways that feed back to affect plant performance. The extent that plant-soil feedback affects co-occurring plant species differentially will determine its impact on plant community structure. Whether feedback operates consistently across similar plant communities is little studied. Here, the same grasses from two eastern U.S. serpentine grasslands and two midwestern tallgrass prairie remnants were examined for plant-soil feedback in parallel greenhouse experiments. Native soils were homogenized and cultured (trained) for a year with each of the four grasses. Feedback was evaluated by examining biomass variation in a second generation of (tester) plants grown in the trained soils. Biomass was lower in soils trained by conspecifics compared to soils trained by heterospecifics in seven of 15 possible cases; biomass was greater in conspecific soils in one other. Sorghastrum nutans exhibited lower biomass in conspecific soils for all four grasslands, so feedback may be characteristic of this species. Three cases from the Hayden prairie site were explained by trainer species having similar effects across all tester species so the relative performance of the different species was little affected; plants were generally larger in soils trained by Andropogon gerardii and smaller in soils trained by S. nutans. Differences among sites in the incidence of feedback were independent of serpentine or prairie soils. To explore the causes of the feedback, several soil factors were measured as a function of trainer species: nutrients and pH, arbuscular mycorrhizal (AM) spore communities, root colonization by AM fungi and putative pathogens, and functional diversity in bacterial communities as indicated by carbon substrate utilization. Only variation in nutrients was consistent with any patterns of feedback, and this could explain the greater biomass in soils trained by A. gerardii at Hayden. Feedback at Nottingham (one of the serpentine sites) differed, most notably for A. gerardii, from that of similar past studies that used different experimental protocols. To understand the consequences of feedback for plant community structure, it is important to consider how multiple species respond to the same plant-induced soil variation as well as differences in the feedback detected between greenhouse and field settings.     

紫茎泽兰入侵过程中生物群落的交互作用

生物入侵在全球范围内影响了生物群落的结构与功能,打破了群落内物种共存的生态格局,继而反馈性影响全球环境。该文就外来杂草紫茎泽兰入侵对生物群落之间交互作用的影响进行了分析。1）紫茎泽兰通过竞争排斥降低了土著植物群落的多样性,造成依赖于土著植物的节肢动物群落减少或丧失适宜的栖息环境。2）打破了土著植物与节肢动物之间相互依存的状态,并通过单优群落优势和强烈化感作用制约天敌昆虫的自然控制作用。3）通过改变地表生境和枯落物种类影响土壤动物群落。4）引起土壤微生物群落组成和功能的变化,改变土壤中可利用资源的形式和数量,影响并重塑了生物种间互作模式,并动态反馈于地面植物群落新格局的形成。分析指出：1）入侵过程中群落之间的交互作用通过多层次生态过程对群落结构与功能的生态改变发挥影响。2）入侵对生物群落的改变所产生的生态驱动反馈性作用于群落互作模式的重塑、群落和生态系统新格局的重建。同时,指出了生物入侵对群落影响的复杂性以及后续研究的方向。     

Plant winners and losers during grassland N-eutrophication differ in biomass allocation and mycorrhizas

Human activities release tremendous amounts of nitrogenous compounds into the atmosphere. Wet and dry deposition distributes this airborne nitrogen (N) on otherwise pristine ecosystems. This eutrophication process significantly alters the species composition of native grasslands; generally a few nitrophilic plant species become dominant while many other species disappear. The functional equilibrium model predicts that, compared to species that decline in response to N enrichment, nitrophilic grass species should respond to N enrichment with greater biomass allocation aboveground and reduced allocation to roots and mycorrhizas. The mycorrhizal feedback hypothesis states that the composition of mycorrhizal fungal communities may influence the composition of plant communities, and it predicts that N enrichment may generate reciprocal shifts in the species composition of mycorrhizal fungi and plants. We tested these hypotheses with experiments that compared biomass allocation and mycorrhizal function of four grass ecotypes (three species), two that gained and two that lost biomass and cover in response to long-term N enrichment experiments at Cedar Creek and Konza Long-Term Ecological Research grasslands. Local grass ecotypes were grown in soil from their respective sites and inoculated with whole-soil inoculum collected from either fertilized (FERT) or unfertilized (UNFERT) plots. Our results strongly support the functional equilibrium model. In both grassland systems the nitrophilic grass species grew taller, allocated more biomass to shoots than to roots, and formed fewer mycorrhizas compared to the grass species that it replaced. Our results did not fully support the hypothesis that N-induced changes in the mycorrhizal fungal community were drivers of the plant community shifts that accompany N eutrophication. The FERT and UNFERT soil inoculum influenced the growth of the grasses differently, but this varied with site and grass ecotype in both expected and unexpected ways suggesting that ambient soil fertility or other factors may be interacting with mycorrhizal feedbacks.     

三江源区不同建植年代人工草地群落演替与土壤养分变化

研究了了三源区不同建植期人工修复草地在不同演替阶段毒杂草[主要是甘肃马先蒿(Pedicularis kansuensis)]的入侵规律、数量特征,植物群落物种组成、生物苗和草地质最以及土壤养分、微生物活性的变化规律.结果表明,不同建植期人工修复草地植物群落的种类组成、植物功能群组成和群落数量特征存在显著差异.随着演替时间的推移,人工草地群落盖度、高度、物种数、生物最和多样性指数均表现出"V"字型变化规律,杂类草--甘肃马先蒿的数量特征变化尤为明显,在4 a的人工草地群落中开始局部入侵,在5～6 a的人工草地群落中大面积入侵,其入侵速度、入侵面积达到高峰期.土壤的含水量、容重、土壤中有机质、氮素和磷素在演替过程(7 a、9 a草地)中逐渐降低,到一定时期又逐步增加;随着演替的进行,不同建植期人工草地的土壤微牛物生物量碳和酶活性均呈"V"字型,变化.对于退化生态系统的恢复首先是植被恢复,其次是土壤肥力的恢复.土壤有机质等养分的积累、微生物活性的改善不仅能使土壤-植物复合系统的功能得以恢复,同时也能促进物种多样性的形成,有利于人工草地群落稳定性的提高.在试验区尽管植被恢复演替进行得比较缓慢,但从土壤发展的角度看,仍属进展演替.所以,在退化高寒草甸的恢复过程中,若降低和有效控制外界的干扰(如围栏封育),可为退化草地恢复提供繁殖体与土壤环境,实现人工草地逐步向恢复(正向)演替进行.图3表6参34     

Evidence for ecological matching of whole AM fungal communities to the local plant-soil environment

The range of ecological roles exhibited by arbuscular mycorrhizal (AM) fungi depends on functional differences among naturally occurring local assemblages of AM species. While functional differences have been demonstrated among AM fungal species and among geographic isolates of the same species, almost nothing is known about functional differences among whole communities of naturally occurring AM fungi. In the greenhouse, we reciprocally transplanted whole AM fungal communities between plant-soil systems representing a serpentine grassland and a tallgrass prairie, using as hosts two grasses common to both systems. For Sorghastrum nutans, native fungi consistently enhanced plant growth more than fungi switched from the alternate system. For Schizachyrium scoparium, foreign and native fungi promoted plant growth similarly in both the serpentine and prairie systems. Thus, the use of foreign inoculum in restoration could change the relative performance, and potentially the competitive abilities, of co-occurring plant species. Moving AM fungal inocula into foreign environments also caused changes in the taxonomic composition of the resultant spore communities, demonstrating their response to environmental influences. These results provide strong evidence for functional differences among naturally occurring AM communities and suggest that a particular AM fungal community may be better matched ecologically to its local habitat than communities taken from other locations.     

Combined effect of fire and water scarcity on vegetation patterns in arid lands

In many arid zones around the word, the vegetation spontaneously forms regular patterns to optimize the use of the scarce water resources. The patterns act as early warning signal that fragile ecosystems may suddenly undergo irreversible shifts, thus, interpreting the structural shape of vegetation patterns is crucial to deciphering the ecosystem history and its expected further development. The sudden and irreversible shift of delicate ecosystems as a consequence of minor variation of the climatic forcing has been studied extensively in the past. The attitude of the ecosystem to recover after a catastrophic event, such as fire, did not receive as much attention so far. Here we modelled fire, as a sudden shift of the ecosystem state variables and functionality and evaluated post-fire scenarios under the hypothesis that two major feedbacks shaped the vegetation patterns: a positive feedback between preferential infiltration and plant growth, and a second feedback between infiltration and vegetation burning. A simple model solving a set of partial differential equations for soil moisture, plant biomass, surface water and dead biomass balance predicted significantly diverse post-fire vegetation patterns depending on the fire severity and on the degree of soil water repellency induced by the vegetation burning.     

Decoupling of component and ensemble density feedbacks in birds and mammals

A component density feedback represents the effect of change in population size on single demographic rates, whereas an ensemble density feedback captures that effect on the overall growth rate of a population. Given that a population's growth rate is a synthesis of the interplay of all demographic rates operating in a population, we test the hypothesis that the strength of ensemble density feedback must augment with increasing strength of component density feedback, using long-term censuses of population size, fertility, and survival rates of 109 bird and mammal populations (97 species). We found that compensatory and depensatory component feedbacks were common (each detected in approximately 50% of the demographic rates). However, component feedback strength only explained <10% of the variation in ensemble feedback strength. To explain why, we illustrate the different sources of decoupling between component and ensemble feedbacks. We argue that the management of anthropogenic impacts on populations using component feedbacks alone is ill-advised, just as managing on the basis of ensemble feedbacks without a mechanistic understanding of the contributions made by its components and environmental variability can lead to suboptimal decisions.     

Additive effects of aboveground polyphagous herbivores and soil feedback in native and range-expanding exotic plants

Plant biomass and plant abundance can be controlled by aboveground and belowground natural enemies. However, little is known about how the aboveground and belowground enemy effects may add up. We exposed 15 plant species to aboveground polyphagous insect herbivores and feedback effects from the soil community alone, as well as in combination. We envisaged three possibilities: additive, synergistic, or antagonistic effects of the aboveground and belowground enemies on plant biomass. In our analysis, we included native and phylogenetically related range-expanding exotic plant species, because exotic plants on average are less sensitive to aboveground herbivores and soil feedback than related natives. Thus, we examined if lower sensitivity of exotic plant species to enemies also alters aboveground-belowground interactions. In a greenhouse experiment, we exposed six exotic and nine native plant species to feedback from their own soil communities, aboveground herbivory by polyphagous insects, or a combination of soil feedback and aboveground insects and compared shoot and root biomass to control plants without aboveground and belowground enemies. We observed that for both native and range-expanding exotic plant species effects of insect herbivory aboveground and soil feedback added up linearly, instead of enforcing or counteracting each other. However, there was no correlation between the strength of aboveground herbivory and soil feedback. We conclude that effects of polyphagous aboveground herbivorous insects and soil feedback add up both in the case of native and related range-expanding exotic plant species, but that aboveground herbivory effects may not necessarily predict the strengths of soil feedback effects.     

Increasing enemy biodiversity strengthens herbivore suppression on two plant species

Concern over biodiversity loss, especially at higher trophic levels, has led to a surge in studies investigating how changes in natural enemy diversity affect community and ecosystem functioning. These studies have found that increasing enemy diversity can strengthen, weaken, and not affect prey suppression, demonstrating that multi-enemy effects on prey are context-dependent. Here we ask how one factor, plant species identity, influences multi-enemy effects on prey. We focused on two plant species of agricultural importance, potato (Solanum tuberosum), and collards (Brassica oleracea L.). These species share a common herbivorous pest, the green peach aphid (Myzus persicae), but vary in structural and chemical traits that affect aphid reproductive rates and which may also influence inter-enemy interactions. In a large-scale field experiment, overall prey exploitation varied dramatically among the plant species, with enemies reducing aphid populations by approximately 94% on potatoes and approximately 62% on collards. Increasing enemy diversity similarly strengthened aphid suppression on both plants, however, and there was no evidence that plant species identity significantly altered the relationship between enemy diversity and prey suppression. Microcosm experiments suggested that, on both collards and potatoes, intraspecific competition among natural enemies exceeded interspecific competition. Enemy species showed consistent and significant differences in where they foraged on the plants, and enemies in the low-diversity treatment tended to spend less time foraging than enemies in the high-diversity treatment. These data suggest that increasing enemy diversity may strengthen aphid suppression because interspecific differences in where enemies forage on the plant allow for greater resource partitioning. Further, these functional benefits of diversity appear to be robust to changes in plant species identity.     

Ecosystem development in roadside grasslands: biotic control, plant-soil interactions, and dispersal limitations

Roadside grasslands undergoing secondary succession are abundant, and represent ecologically meaningful examples of novel, human-created ecosystems. Interactions between plant and soil communities (hereafter plant-soil interactions) are of major importance in understanding the role of biotic control in ecosystem functioning, but little is known about these links in the context of ecosystem restoration and succession. The assessment of the key biotic communities and interactions driving ecosystem development will help practitioners to better allocate the limited resources devoted to roadside grassland restoration. We surveyed roadside grasslands from three successional stages (0-2, 7-9, and >20 years) in two Mediterranean regions of Spain. Structural equation modeling was used to evaluate how interactions between plants, biological soil crusts (BSCs), and soil microbial functional diversity (soil microorganisms) affect indicators of ecosystem development and restoration: plant similarity to the reference ecosystem, erosion control, and soil C storage and N accumulation. Changes in plant community composition along the successional gradient exerted the strongest influence on these indicators. High BSC cover was associated with high soil stability, and high soil microbial functional diversity from late-successional stages was associated with high soil fertility. Contrary to our expectations, the indirect effects of plants, mediated by either BSCs or soil microorganisms, were very weak in both regions, suggesting a minor role for plant-soil interactions upon ecosystem development indicators over long periods. Our results suggest that natural vegetation dynamics effectively improved ecosystem development within a time frame of 20 years in the grasslands evaluated. They also indicate that this time could be shortened if management actions focus on: (1) maintaining well-conserved natural areas close to roadsides to enhance plant compositional changes towards late-successional stages, (2) increasing BSC cover in areas under strong erosion risk, to avoid soil loss, and (3) enhancing soil microbial functional diversity in resource-limited areas, to enhance soil C and N accumulation.     

Species Migrations and Ecosystem Stability During Climate Change: The Belowground Connection

Abstract: Compatibility between the belowground mutualists of resident species and the needs of immigrant species will strongly influence the successful transition from one perennial plant community to another during climate change. A hiatus in the overlap between plant species that maintain a positive link with the soil ecosystem could result in site capture by weeds and rapid degradation of the productive capacity of soils. We discuss instances in which such rapid degradation has occurred and argue for the crucial importance of protecting plant-soil links in the coming decades through maintaining biodiversity and utilizing management practices that help plants keep a firm grip on the soil. Examples of the latter include partial and dispersed cutting in forestry, use of green cover crops in agriculture and grazing intensities that permit degraded range to rebuild.     

Compositional stability and diversity of vascular plant communities following logging disturbance in Appalachian forests

Human-caused changes in disturbance regimes and introductions of nonnative species have the potential to result in widespread, directional changes in forest community structure. The degree that plant community composition persists or changes following disturbances depends on the balance between local extirpation and colonization by new species, including nonnatives. In this study, we examined species losses and gains, and entry of native vs. exotic species to determine how oak forests in the Appalachian Mountains might shift in species composition following a gradient of pulse disturbances (timber harvesting). We asked (1) how compositional stability of the plant community (resistance and resilience) was influenced by disturbance intensity, (2) whether community responses were driven by extirpation or colonization of species, and (3) how disturbance intensity influenced total and functional group diversity, including the nonnative proportion of the flora through time. We collected data at three spatial scales and three times, including just before, one year post-disturbance, and 10 years post-disturbance. Resistance was estimated using community distance measures between pre- and one year post-disturbance, and resilience using community distance between pre- and 10-year post-disturbance conditions. The number of colonizing and extirpated species between sampling times was analyzed for all species combined and for six functional groups. Resistance and resilience decreased with increasing timber-harvesting disturbance; compositional stability was lower in the most disturbed plots, which was driven by colonization, but not extirpation, of species. Colonization of species also led to increases in diversity after disturbance that was typically maintained after 10 years following disturbance. Most of the community-level responses were driven by post-disturbance colonization of native forbs and graminoids. The nonnative proportion of plant species tended to increase following disturbance, especially at large spatial scales in the most disturbed treatments, but tended to decrease through time following disturbance due to canopy development. The results of this study are consistent with the theory that resources released by disturbance have strong influences on species colonization and community composition. The effects of management activities tested in this study, which span a gradient of timber-harvesting disturbance, shift species composition largely via an increase in species colonization and diversity.     

Decomposers (Lumbricidae, Collembola) affect plant performance in model grasslands of different diversity

Decomposer invertebrates influence soil structure and nutrient mineralization as well as the activity and composition of the microbial community in soil and therefore likely affect plant performance and plant competition. We established model grassland communities in a greenhouse to study the interrelationship between two different functional groups of decomposer invertebrates, Lumbricidae and Collembola, and their effect on plant performance and plant nitrogen uptake in a plant diversity gradient. Common plant species of Central European Arrhenatherion grasslands were transplanted into microcosms with numbers of plant species varying from one to eight and plant functional groups varying from one to four. Separate and combined treatments with earthworms and collembolans were set up. Microcosms contained 15N labeled litter to track N fluxes into plant shoots. Presence of decomposers strongly increased total plant and plant shoot biomass. Root biomass decreased in the presence of collembolans and even more in the presence of earthworms. However, it increased when both animal groups were present. Also, presence of decomposers increased total N concentration and 15N enrichment of grasses, legumes, and small herbs. Small herbs were at a maximum in the combined treatment with earthworms and collembolans. The impact of earthworms and collembolans on plant performance strongly varied with plant functional group identity and plant species diversity and was modified when both decomposers were present. Both decomposer groups generally increased aboveground plant productivity through effects on litter decomposition and nutrient mineralization leading to an increased plant nutrient acquisition. The non-uniform effects of earthworms and collembolans suggest that functional diversity of soil decomposer animals matters and that the interactions between soil animal functional groups affect the structure of plant communities.     

Native plant diversity resists invasion at both low and high resource levels

Human modification of the environment is causing both loss of species and changes in resource availability. While studies have examined how species loss at the local level can influence invasion resistance, interactions between species loss and other components of environmental change remain poorly studied. In particular, the manner in which native diversity interacts with resource availability to influence invasion resistance is not well understood. We created experimental plant assemblages that varied in native species (1-16 species) and/or functional richness (defined by rooting morphology and phenology; one to five functional groups). We crossed these diversity treatments with resource (water) addition to determine their interactive effects on invasion resistance to spotted knapweed (Centaurea maculosa), a potent exotic invader in the intermountain West of the United States. We also determined how native diversity and resource addition influenced plant-available soil nitrogen, soil moisture, and light. Assemblages with lower species and functional diversity were more heavily invaded than assemblages with greater species and functional diversity. In uninvaded assemblages, experimental addition of water increased soil moisture and plant-available nitrogen and decreased light availability. The availability of these resources generally declined with increasing native plant diversity. Although water addition increased susceptibility to invasion, it did not fundamentally change the negative relationship between diversity and invasibility. Thus, native diversity provided strong invasion resistance even under high resource availability. These results suggest that the effects of local diversity can remain robust despite enhanced resource levels that are predicted under scenarios of global change.     

Dispersal, competition, and shifting patterns of diversity in a degraded oak savanna

Diversity is a balance between processes that add and limit species (e.g., dispersal vs. competition), but reconciling their contributions remains a challenge. Recruit-ment-based models predict that dispersal barriers are most limiting for diversity, while competition-based models predict that dispersal matters only when competition is minimized. Testing these models is difficult because their influence varies with scale and site productivity. In a degraded oak savanna, we used plot-level (seed additions, burning) and site-level (proportions of regional functional groups found locally) analyses in areas with variable soil depth to examine how dispersal and competition influence diversity. At the plot level, added species persisted where they were formerly absent, but few established naturally despite fire-induced resource enrichment and nearby populations, revealing the importance of dispersal limitation for diversity. This result did not vary with soil depth or standing crop. Although competition could not prevent establishment in unburned plots, it significantly lowered survival, indicating that resource limitations exacerbate dispersal inefficiencies. At the site level, the concordance between regional and local diversity for native species was associated with soil depth heterogeneity, not dispersal or competition. This suggests that persistence is determined primarily by the influence of the environment on population demographics. Given that the formation of new populations is unlikely, those remaining appear to be confined to optimal habitat where they resist competitive or stochastic displacement, possibly explaining why species loss is rare despite substantial habitat loss and invasion. For exotics, there was no relationship between diversity and soil depth heterogeneity. Annuals with presumed dispersal capabilities were significantly overrepresented in all sites while perennial forbs, the largest regional functional group, were significantly underrepresented. We interpret the native-exotic discrepancies as reflecting the recent arrival of exotics (150 years ago), suggesting that local establishment occurs slowly even for species with regional prevalence. The accumulation lag may be explained by the need for founder populations to be demographically stable; otherwise persistence requires continual immigration favoring overrepresentation by dispersers. Our findings support the view that dispersal limitation restricts diversity within plant communities, but suggests that the impacts of environment on demographic performance ultimately determine the pattern and rate of community assembly.     

Soil feedback of exotic savanna grass relates to pathogen absence and mycorrhizal selectivity

Enemy release of exotic plants from soil pathogens has been tested by examining plant-soil feedback effects in repetitive growth cycles. However, positive soil feedback may also be due to enhanced benefit from the local arbuscular mycorrhizal fungi (AMF). Few studies actually have tested pathogen effects, and none of them did so in arid savannas. In the Kalahari savanna in Botswana, we compared the soil feedback of the exotic grass Cenchrus biflorus with that of two dominant native grasses, Eragrostis lehmanniana and Aristida meridionalis. The exotic grass had neutral to positive soil feedback, whereas both native grasses showed neutral to negative feedback effects. Isolation and testing of root-inhabiting fungi of E. lehmanniana yielded two host-specific pathogens that did not influence the exotic C. biflorus or the other native grass, A. meridionalis. None of the grasses was affected by the fungi that were isolated from the roots of the exotic C. biflorus. We isolated and compared the AMF community of the native and exotic grasses by polymerase chain reaction-denaturing gradient gel elecrophoresis (PCR-DGGE), targeting AMF 18S rRNA. We used roots from monospecific field stands and from plants grown in pots with mixtures of soils from the monospecific field stands. Three-quarters of the root samples of the exotic grass had two nearly identical sequences, showing 99% similarity with Glomus versiforme. The two native grasses were also associated with distinct bands, but each of these bands occurred in only a fraction of the root samples. The native grasses contained a higher diversity of AMF bands than the exotic grass. Canonical correspondence analyses of the AMF band patterns revealed almost as much difference between the native and exotic grasses as between the native grasses. In conclusion, our results support the hypothesis that release from soil-borne enemies may facilitate local abundance of exotic plants, and we provide the first evidence that these processes may occur in arid savanna ecosystems. Pathogenicity tests implicated the involvement of soil pathogens in the soil feedback responses, and further studies should reveal the functional consequences of the observed high infection with a low diversity of AMF in the roots of exotic plants.