Embracing variability: environmental dependence and plant community context in ecological restoration

In heterogeneous landscapes, one can expect a complexity of ecological restoration outcomes. The effectiveness of management often depends on environmental conditions (environmental context) and how management indirectly affects other components of the system (community context). Although managers appreciate this context dependency, it is difficult to translate it to decision‐making in restoration. We demonstrate one approach to improve this translation. We surveyed plant, soil, and landscape characteristics at 131 grassland and coastal sage sites that received herbicide treatments to remove non‐native plant species and/or propagule addition to increase native species. We used path analysis to describe how each management approach influenced target non‐native species and how interactions with environmental conditions and indirect effects of management influenced plant community composition. This approach enabled us to analyze a complex system with differing management histories to identify both direct and indirect effects of management. Management had the intended direct effects: the application of herbicide and propagule addition directly reduced non‐native species and increased native species, respectively. We found little evidence of environmental dependency: effects occurred largely independent of environmental conditions. However, management outcomes did depend on plant community context. Specifically, although herbicide reduced the cover of target, non‐native plant species, this reduction resulted in only slight increases in native species and instead led indirectly to increases in non‐targeted, non‐native species. We suggest that quantitative evaluation of variability in restoration outcomes allows management to be more adaptive and increase decision‐making efficacy in complex managed landscapes.     

Soil Organism and Plant Introductions in Restoration of Species-Rich Grassland Communities

Soil organisms can strongly affect competitive interactions and successional replacements of grassland plant species. However, introduction of whole soil communities as management strategy in grassland restoration has received little experimental testing. In a 5-year field experiment at a topsoil-removed ex-arable site ( receptor site ), we tested effects of (1) spreading hay and soil, independently or combined, and (2) transplanting intact turfs on plant and soil nematode community development. Material for the treatments was obtained from later successional, species-rich grassland ( donor site ). Spreading hay affected plant community composition, whereas spreading soil did not have additional effects. Plant species composition of transplanted turfs became less similar to that in the donor site. Moreover, most plants did not expand into the receiving plots. Soil spreading and turf transplantation did not affect soil nematode community composition. Unfavorable soil conditions (e.g., low organic matter content and seasonal fluctuations in water level) at the receptor site may have limited plant and nematode survival in the turfs and may have precluded successful establishment outside the turfs. We conclude that introduction of later successional soil organisms into a topsoil-removed soil did not facilitate the establishment of later successional plants, probably because of the "mismatch" in abiotic soil conditions between the donor and the receptor site. Further research should focus on the required conditions for establishment of soil organisms at restoration sites in order to make use of their contribution to grassland restoration. We propose that introduction of organisms from "intermediate" stages will be more effective as management strategy than introduction of organisms from "target" stages.     

Single introductions of soil biota and plants generate long‐term legacies in soil and plant community assembly

Recent demonstrations of the role of plant–soil biota interactions have challenged the conventional view that vegetation changes are mainly driven by changing abiotic conditions. However, while this concept has been validated under natural conditions, our understanding of the long‐term consequences of plant–soil interactions for above‐belowground community assembly is restricted to mathematical and conceptual model projections. Here, we demonstrate experimentally that one‐time additions of soil biota and plant seeds alter soil‐borne nematode and plant community composition in semi‐natural grassland for 20 years. Over time, aboveground and belowground community composition became increasingly correlated, suggesting an increasing connectedness of soil biota and plants. We conclude that the initial composition of not only plant communities, but also soil communities has a long‐lasting impact on the trajectory of community assembly.     

Site preparation impacts on soil biotic and abiotic properties,weed control,and native grass establishment

In severely degraded systems active restoration is required to overcome legacies of past land use and to create conditions that promote the establishment of target plant communities. While our understanding of the importance of soil microbial communities in ecological restoration is growing, few studies have looked at the impacts different site preparation techniques have on these communities. We trialed four methods of site preparation: fire, top‐soil removal (TSR; removal of top 50 mm of soil), slashing (vegetation cut to 30 mm, biomass removed), and carbon (C; as sugar and saw‐dust) addition, and quantified resulting soil bacterial communities using DNA metabarcoding. We compared the effectiveness of these techniques to reduce weed biomass, improve native grass establishment, and induce changes in soil nutrient availability. TSR was the most effective technique, leading to a reduction in both available nutrients and competition from weeds. In comparison, the remaining methods had little or no effect on weed biomass, native grass establishment, or soil nutrient availability. Both TSR and C addition resulted in changes in the soil bacterial community. These changes have the potential to alter plant community assembly in many ways, such as via nutrient acquisition, pathogenic effects, nutrient cycling, and decomposition. We recommend TSR for ecological restoration of old‐fields and suggest it is a much more effective technique than burning, slashing, or C addition. Restoration practitioners should consider how their management techniques may influence the soil biota and, in turn, affect restoration outcomes.     

Soil acidification reduces the effects of short‐term nutrient enrichment on plant and soil biota and their interactions in grasslands

Soil nitrogen (N) and phosphorus (P) contents, and soil acidification have greatly increased in grassland ecosystems due to increased industrial and agricultural activities. As major environmental and economic concerns worldwide, nutrient enrichment and soil acidification can lead to substantial changes in the diversity and structure of plant and soil communities. Although the separate effects of N and P enrichment on soil food webs have been assessed across different ecosystems, the combined effects of N and P enrichment on multiple trophic levels in soil food webs have not been studied in semiarid grasslands experiencing soil acidification. Here we conducted a short‐term N and P enrichment experiment in non‐acidified and acidified soil in a semiarid grassland on the Mongolian Plateau. We found that net primary productivity was not affected by N or P enrichment alone in either non‐acidified or acidified soil, but was increased by combined N and P enrichment in both non‐acidified and acidified soil. Nutrient enrichment decreased the biomass of most microbial groups in non‐acidified soil (the decrease tended to be greatest with combined N and P enrichment) but not in acidified soil, and did not affect most soil nematode variables in non‐acidified or acidified soil. Nutrient enrichment also changed plant and microbial community structure in non‐acidified but not in acidified soil, and had no effect on nematode community structure in non‐acidified or acidified soil. These results indicate that the responses to short‐term nutrient enrichment were weaker for higher trophic groups (nematodes) than for lower trophic groups (microorganisms) and primary producers (plants). The findings increase our understanding of the effects of nutrient enrichment on multiple trophic levels of soil food webs, and highlight that soil acidification, as an anthropogenic stressor, reduced the responses of plants and soil food webs to nutrient enrichment and weakened plant–soil interactions.     

Soil-mediated indirect impacts of an invasive predator on plant growth

While several studies have shown that invasive plant effects on soil biota influence subsequent plant performance, corresponding studies on how invasive animals affect plants through influencing soil biota are lacking. This is despite the fact that invasive animals often indirectly alter the below-ground subsystem. We studied 18 offshore islands in northern New Zealand, half of which have been invaded by rats that are predators of seabirds and severely reduce their densities, and half of which remain non-invaded; invasion of rats thwarts seabird transfer of resources from ocean to land. We used soil from each island in a glasshouse experiment involving soil sterilization treatments to determine whether rat invasion indirectly influences plant growth through the abiotic pathway (by impairing seabird-driven inputs to soil) or the biotic pathway (by altering the soil community). Rat invasion greatly impaired plant growth but entirely through the abiotic pathway. Plant growth was unaffected by the soil community or its response to invasion, meaning that the responses of plants and soil biota to invasion are decoupled. Our results provide experimental evidence for the powerful indirect effects that predator-instigated cascades can exert on plant and ecosystem productivity, with implications for the restoration of island ecosystems by predator removal.     

Diverse responses across soil parent materials during ecological restoration

A major challenge to advancing the science and practice of ecological restoration is working across large landscapes containing diverse sites that may respond differently to restoration. We conducted a 5‐year restoration experiment, replicated across 9 sites spanning 3 soil parent material types within a 9,000‐ha Pinus ponderosa forest landscape. We evaluated plant community response to restoration Pinus thinning, grazing, and aqueous smoke application. We measured vegetation before (2003) and 3 (2006) and 5 (2008) years after treatment. Plant community responses of species richness, cover, and composition were diverse, ranging from increases, decreases, or no change depending on soil parent material, tree thinning, and presence or exclusion of grazing. Restoration outcomes were under hierarchical control: soil parent material constrained response to Pinus thinning, which in turn influenced grazing effects. On limestone‐derived soil, responses included no change in species richness but increased plant cover with Pinus thinning. Both plant richness and cover increased on benmorite soil after thinning, and cover generally increased more without grazing. On rocky, basalt soil, plant richness increased but cover did not after any treatment. Diversity of responses to restoration has implications for: (1) setting goals or monitoring indicators tailored to inherent soil capability; (2) identifying where grazing most affects restoration outcomes; and (3) forecasting responses to restoration across landscapes. Diverse responses to restoration along physiographic gradients such as soil parent material warrant consideration when developing restoration across degraded landscapes.     

Genetic by environment interactions affect plant–soil linkages

The role of plant intraspecific variation in plant–soil linkages is poorly understood, especially in the context of natural environmental variation, but has important implications in evolutionary ecology. We utilized three 18‐ to 21‐year‐old common gardens across an elevational gradient, planted with replicates of five Populus angustifolia genotypes each, to address the hypothesis that tree genotype (G), environment (E), and G × E interactions would affect soil carbon and nitrogen dynamics beneath individual trees. We found that soil nitrogen and carbon varied by over 50% and 62%, respectively, across all common garden environments. We found that plant leaf litter (but not root) traits vary by genotype and environment while soil nutrient pools demonstrated genotype, environment, and sometimes G × E interactions, while process rates (net N mineralization and net nitrification) demonstrated G × E interactions. Plasticity in tree growth and litter chemistry was significantly related to the variation in soil nutrient pools and processes across environments, reflecting tight plant–soil linkages. These data overall suggest that plant genetic variation can have differential affects on carbon storage and nitrogen cycling, with implications for understanding the role of genetic variation in plant–soil feedback as well as management plans for conservation and restoration of forest habitats with a changing climate.     

Trophic and non‐trophic interactions influence the mechanisms underlying biodiversity–ecosystem functioning relationships under different abiotic conditions

Plant diversity effects on ecosystem functioning usually have been studied from a plant perspective. However, the mechanisms underlying biodiversity–ecosystem functioning relationships may also depend on positive or negative interactions between plants and other biotic and abiotic factors, which remain poorly understood. Here we assessed whether plant–herbivore and/or plant–detritivore interactions modify the biodiversity–ecosystem functioning relationship and the mechanisms underlying biodiversity effects, including complementarity and selection effects, biomass allocation, vertical distribution of roots, and plant survival using a microcosm experiment. We also evaluated to what extent trophic and non‐trophic interactions are affected by abiotic conditions by studying drought effects. Our results show that biotic and abiotic conditions influence the shape of the biodiversity–ecosystem function relationship, varying from hump‐shaped to linear. For instance, total biomass increased linearly with plant richness in the presence of detritivores, but not in the absence of detritivores. Moreover, detritivore effects on belowground plant productivity were highly context dependent, varying in the presence of herbivores. Plant interactions with soil biota, especially with herbivores, influenced the mechanisms underlying diversity effects. Herbivores increased plant complementarity and modified biomass allocation and vertical distribution of roots. Furthermore, biotic–abiotic interactions influenced plant productivity differently across plant functional groups. Our findings emphasize the importance of complex biotic interactions underlying biodiversity effects, and that these biotic interactions may change with abiotic conditions. Despite minor changes in productivity in the short‐term, soil biota‐induced changes in plant–plant interactions and plant survival are likely to have significant long‐term consequences for ecosystem functioning. Considering the context‐dependency of multichannel interactions may contribute to reconciling differences among observed patterns in biodiversity studies. Further, abiotic conditions modified the effects of biotic interactions, suggesting that changes in environmental conditions may not only affect ecosystems directly, but also change the biotic composition of and dynamics within ecosystems.     

Resilience and restoration of tropical and subtropical grasslands,savannas, and grassy woodlands

Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open‐canopy grassy woodlands) remains limited. To incorporate grasslands into large‐scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved – frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human‐caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species‐diverse plant communities, including endemic species, are slow to recover. Complicating plant‐community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re‐establish. To guide restoration decisions, we draw on the old‐growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old‐growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old‐growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.     

Interactions between soil heterogeneity and freezing: Implications for grassland plant diversity and relative species abundances

Plant stress resulting from soil freezing is expected to increase in northern temperate regions over the next century due to reductions in snow cover caused by climate change. Within plant communities, soil spatial heterogeneity can potentially buffer the effects of plant freezing stress by increasing the availability of soil microsites that function as microrefugia. Moreover, increased species richness resulting from soil heterogeneity can increase the likelihood of stress‐tolerant species being present in a community. We used a field experiment to examine interactions between soil heterogeneity and increased freezing intensity (achieved via snow removal) on plant abundance and diversity in a grassland. Patches of topsoil were mixed with either sand or woodchips to create heterogeneous and homogeneous treatments, and plant community responses to snow removal were assessed over three growing seasons. Soil heterogeneity interacted significantly with snow removal, but it either buffered or exacerbated the snow removal response depending on the specific substrate (sand vs. woodchips) and plant functional group. In turn, snow removal influenced plant responses to soil heterogeneity; for example, adventive forb cover responded to increased heterogeneity under ambient snow cover, but this effect diminished with snow removal. Our results reveal that soil heterogeneity can play an important role in determining plant responses to changes in soil freezing stress resulting from global climate change. While the deliberate creation of soil microsites in ecological restoration projects as a land management practice could increase the frequency of microrefugia that mitigate plant community responses to increased freezing stress, the design of these microsites must be optimized, given that soil heterogeneity also has the potential to exacerbate freezing stress responses.     

How does plant richness affect pollinator richness and temporal stability of flower visits?

Pollinators play a key role in the reproduction of most plant species, and pollinator and plant diversity are often related. We studied an experimental gradient of plant species richness for a better understanding of plant–pollinator community interactions and their temporal variability, because in non‐experimental field surveys plant richness is often confounded with gradients in management, soil fertility, and community composition. We observed pollinator species richness and frequency of visits six times in 73 plots over two years, and used advanced statistical analysis to account for the high number of zeroes that often occur in count data of rare species. The frequency of pollinator visits increased linearly with both the blossom cover and the number of flowering plant species, which was closely related to the total number of plant species, whereas the number of pollinator species followed a saturation curve. The presence of particularly attractive plant species was only important for the frequency of flower visits, but not to the richness of pollinators. Plant species richness, blossom cover, and the presence of attractive plant species enhanced the temporal stability in the frequency of pollinator visits. In conclusion, grasslands with high plant diversity enhance and stabilize frequent and diverse flower visitations, which should sustain effective pollination and plant reproduction.     

Winning and losing with microbes: how microbially mediated fitness differences influence plant diversity

Interactions between plants and soil microbes can strongly influence plant diversity and community dynamics. Soil microbes may promote plant diversity by driving negative frequency‐dependent plant population dynamics, or may favor species exclusion by providing one species an average fitness advantage over others. However, past empirical research has focused overwhelmingly on the consequences of frequency‐dependent feedbacks for plant species coexistence and has generally neglected the consequences of microbially mediated average fitness differences. Here we use theory to develop metrics that quantify microbially mediated plant fitness differences, and show that accounting for these effects can profoundly change our understanding of how microbes influence plant diversity. We show that soil microbes can generate fitness differences that favour plant species exclusion when they disproportionately harm (or favour) one plant species over another, but these fitness differences may also favor coexistence if they trade off with competition for other resources or generate intransitive dominance hierarchies among plants. We also show how the metrics we present can quantify microbially mediated fitness differences in empirical studies, and explore how microbial control over coexistence varies along productivity gradients. In all, our analysis provides a more complete theoretical foundation for understanding how plant–microbe interactions influence plant diversity.     

Plant‐driven changes in soil microbial communities influence seed germination through negative feedbacks

相似文献   

Diversity and salt tolerance of native Acacia rhizobia isolated from saline and non‐saline soils

Re‐establishing native vegetation in stressed soils is of considerable importance in many parts of the world, leading to significant interest in using plant–soil symbiont interactions to increase the cost‐effectiveness of large‐scale restoration. However, effective use of soil microbes in revegetation requires knowledge of how microbe communities vary along environmental stress gradients, as well as how such variation relates to symbiont effectiveness. In Australia, shrubby legumes dominate many ecosystems where dryland salinity is a major issue, and improving plant establishment in saline soils is a priority of regional management agencies. In this study, strains of rhizobial bacteria were isolated from a range of Acacia spp. growing in saline and non‐saline soils. Replicates of each strain were grown under several salinity levels in liquid culture and characterized for growth and salt tolerance. Genetic characterization of rhizobia showed considerable variation among strains, with salt tolerance and growth generally higher in rhizobial populations derived from more saline soils. These strains showed markedly different genetic profiles and generic affiliations to those from more temperate soils, suggesting community differentiation in relation to salt stress. The identification of novel genomic species from saline soils suggests that the diversity of rhizobia associated with Australian Acacia spp. is significantly greater than previously described. Overall, the ability of some symbiotically effective strains to tolerate high salinity is promising with regard to improving host plant re‐establishment in these soils.     

Context‐dependent biotic interactions control plant abundance across altitudinal environmental gradients

Many biotic interactions influence community structure, yet most distribution models for plants have focused on plant competition or used only abiotic variables to predict plant abundance. Furthermore, biotic interactions are commonly context‐dependent across abiotic gradients. For example, plant–plant interactions can grade from competition to facilitation over temperature gradients. We used a hierarchical Bayesian framework to predict the abundances of 12 plant species across a mountain landscape and test hypotheses on the context‐dependency of biotic interactions over abiotic gradients. We combined field‐based estimates of six biotic interactions (foliar herbivory and pathogen damage, fungal root colonization, fossorial mammal disturbance, plant cover and plant diversity) with abiotic data on climate and soil depth, nutrients and moisture. All biotic interactions were significantly context‐dependent along temperature gradients. Results supported the stress gradient hypothesis: as abiotic stress increased, the strength or direction of the relationship between biotic variables and plant abundance generally switched from negative (suggesting suppressed plant abundance) to positive (suggesting facilitation/mutualism). For half of the species, plant cover was the best predictor of abundance, suggesting that the prior focus on plant–plant interactions is well‐justified. Explicitly incorporating the context‐dependency of biotic interactions generated novel hypotheses about drivers of plant abundance across abiotic gradients and may improve the accuracy of niche models.     

Soil legacy effects of plants and drought on aboveground insects in native and range-expanding plant communities

Soils contain biotic and abiotic legacies of previous conditions that may influence plant community biomass and associated aboveground biodiversity. However, little is known about the relative strengths and interactions of the various belowground legacies on aboveground plant–insect interactions. We used an outdoor mesocosm experiment to investigate the belowground legacy effects of range-expanding versus native plants, extreme drought and their interactions on plants, aphids and pollinators. We show that plant biomass was influenced more strongly by the previous plant community than by the previous summer drought. Plant communities consisted of four congeneric pairs of natives and range expanders, and their responses were not unanimous. Legacy effects affected the abundance of aphids more strongly than pollinators. We conclude that legacies can be contained as soil ‘memories’ that influence aboveground plant community interactions in the next growing season. These soil-borne ‘memories’ can be altered by climate warming-induced plant range shifts and extreme drought.     

When and where plant‐soil feedback may promote plant coexistence: a meta‐analysis

Plant‐soil feedback (PSF) theory provides a powerful framework for understanding plant dynamics by integrating growth assays into predictions of whether soil communities stabilise plant–plant interactions. However, we lack a comprehensive view of the likelihood of feedback‐driven coexistence, partly because of a failure to analyse pairwise PSF, the metric directly linked to plant species coexistence. Here, we determine the relative importance of plant evolutionary history, traits, and environmental factors for coexistence through PSF using a meta‐analysis of 1038 pairwise PSF measures. Consistent with eco‐evolutionary predictions, feedback is more likely to mediate coexistence for pairs of plant species (1) associating with similar guilds of mycorrhizal fungi, (2) of increasing phylogenetic distance, and (3) interacting with native microbes. We also found evidence for a primary role of pathogens in feedback‐mediated coexistence. By combining results over several independent studies, our results confirm that PSF may play a key role in plant species coexistence, species invasion, and the phylogenetic diversification of plant communities.     

Vegetation succession on reclaimed coal wastes in Spain: the influence of soil and environmental factors

Question: How is vegetation succession on coal mine wastes under a Mediterranean climate affected by the restoration method used (topsoil addition or not)? How are plant successional processes influenced by local landscape and soil factors? Location: Reclaimed coal mines in the north of Palencia province, northern Spain (42°47′‐42°50′ N, 4°32′‐4°53′ W). Methods: In Jun–Jul 2008, vascular plant species cover was monitored in 31 coal mines. The mines, which had been restored using two restoration methods (topsoil addition or not), comprised a chronosequence of different ages from 1 to 40 yr since restoration started. Soil and environmental factors at each mine were monitored and related to species cover using a combination of ordination methods and Huisman–Olff–Fresco modeling. Results: Plant succession was affected by restoration method . Where topsoil was added, succession was influenced by age since restoration and soil pH. Where no topsoil was added, soil factors seem to arrest succession. Vegetation composition on topsoiled sites showed a gradient with age, from the youngest, with early colonizing species, to oldest, with an increase in woody species. Vegetation on non‐topsoiled sites comprised mainly early‐successional species. Response to age and pH of 37 species found on topsoiled mines is described. Conclusions: Restoration of coal mines under this Mediterranean climate can be relatively fast if topsoil is added, with a native shrub community developing after 15 yr. However, if topsoil is not used, it takes more than 40 yr. For topsoiled mines, the species found in the different successional stages were identified, and their tolerance to soil pH was derived. This information will assist future restoration projects in the area.     

Native and non‐native grasses generate common types of plant–soil feedbacks by altering soil nutrients and microbial communities

Soil conditioning occurs when plants alter features of their soil environment. When these alterations affect subsequent plant growth, it is a plant soil feedback. Plant–soil feedbacks are an important and understudied aspect of aboveground–belowground linkages in plant ecology that influence plant coexistence, invasion and restoration. Here, we examine plant–soil feedback dynamics of seven co‐occurring native and non‐native grass species to address the questions of how plants modify their soil environment, do those modifications inhibit or favor their own species relative to other species, and do non‐natives exhibit different plant–soil feedback dynamics than natives. We used a two‐phase design, wherein a first generation of plants was grown to induce species‐specific changes in the soil and a second generation of plants was used as a bioassay to determine the effects of those changes. We also used path‐analysis to examine the potential chain of effects of the first generation on soil nutrients and soil microbial composition and on bioassay plant performance. Our findings show species‐specific (rather than consistent within groups of natives and non‐natives) soil conditioning effects on both soil nutrients and the soil microbial community by plants. Additionally, native species produced plant–soil feedback types that benefit other species more than themselves and non‐native invasive species tended to produce plant–soil feedback types that benefit themselves more than other species. These results, coupled with previous field observations, support hypotheses that plant–soil feedbacks may be a mechanism by which some non‐native species increase their invasive potential and plant–soil feedbacks may influence the vulnerability of a site to invasion.