Plant invasion alters nitrogen cycling by modifying the soil nitrifying community

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

Soil microbial communities adapt to genetic variation in leaf litter inputs

Plant genotypes can have important community‐ and ecosystem‐level effects. However, whether the extended phenotypes of plants feed back to influence the fitness of causal genotypes through soil processes remains unknown. We investigated whether aspen genotypes create distinct soil microbial communities that could potentially affect plant fitness. Using naturally occurring aspen stands in an old‐field system, we set up reciprocal litter transplants among ten genetically distinct aspen clones and tracked decomposition and changes in belowground nutrients and microbial communities for three years. We found that belowground microbial communities became adapted to process specific genotypes of aspen litter to the extent allowable by environment and litter chemistry. Belowground processes were driven by a combination of little quality and prior exposure to specific genotypes of litter. In general, litter from aspen genotypes native to the soil community decomposed more rapidly than did litter from foreign aspen genotypes (i.e. a home‐field advantage existed). While home‐field advantages have been documented to occur among litters of different species, we show that intraspecific variation can elicit similar, albeit weak, effects within a single species. Because rapid decomposition and nutrient cycling is likely to benefit fast‐growing, early‐successional species such as aspen, genotype‐mediated selection for soil microbial communities may feed back to positively affect plant fitness. In addition, belowground communities exhibited significant shifts in response to leaf litter inputs. When exposed to foreign litter, microbial communities changed to become more similar to the microbial community beneath the foreign litter's origin, indicating that belowground microbial communities are predictable given the genotype of the aboveground aspen clone.     

Effects of experimental harvesting of an invasive hybrid cattail on wetland structure and function

Invasive plants, such as the hybrid cattail Typha × glauca, can reduce biodiversity and alter the ability of wetlands to provide critical ecosystem services, including nutrient cycling and carbon storage. Several approaches have been used to reduce Typha dominance and restore invaded wetlands, but long‐term studies assessing benefits of these restoration efforts are limited. A previous study demonstrated that aboveground harvesting of Typha × glauca stems and litter reduced Typha dominance 2 years post‐treatment in a Great Lakes coastal wetland. In the current study, we extended monitoring of experimental aboveground Typha harvest to 4 years post‐treatment and added assessments of treatment effects on soil nutrients, carbon emissions, and microbial community composition. Aboveground harvest treatment resulted in a dramatic reduction in Typha litter cover that persisted for 4 years, increased soil temperature, and increased abundance of the native plant genus Carex. However, aboveground harvest treatment did not significantly reduce Typha abundance, nor did it have significant effects on soil nutrient concentrations, carbon fluxes, or the taxonomic composition of soil microbial communities. We did observe differences in bacterial community composition between plots based on time since Typha invasion, which may indicate some legacy effects of Typha invasion. At the scale of this experiment (4 × 4 m plots), our results indicate that a single aboveground removal of Typha × glauca is not sufficient to restore a heavily invaded freshwater wetland ecosystem, and that periodic harvesting of Typha stems and litter may be required to maintain native plant abundance.     

Experimental litterfall manipulation drives large and rapid changes in soil carbon cycling in a wet tropical forest

Global changes such as variations in plant net primary production are likely to drive shifts in leaf litterfall inputs to forest soils, but the effects of such changes on soil carbon (C) cycling and storage remain largely unknown, especially in C‐rich tropical forest ecosystems. We initiated a leaf litterfall manipulation experiment in a tropical rain forest in Costa Rica to test the sensitivity of surface soil C pools and fluxes to different litter inputs. After only 2 years of treatment, doubling litterfall inputs increased surface soil C concentrations by 31%, removing litter from the forest floor drove a 26% reduction over the same time period, and these changes in soil C concentrations were associated with variations in dissolved organic matter fluxes, fine root biomass, microbial biomass, soil moisture, and nutrient fluxes. However, the litter manipulations had only small effects on soil organic C (SOC) chemistry, suggesting that changes in C cycling, nutrient cycling, and microbial processes in response to litter manipulation reflect shifts in the quantity rather than quality of SOC. The manipulation also affected soil CO ₂ fluxes; the relative decline in CO ₂ production was greater in the litter removal plots (?22%) than the increase in the litter addition plots (+15%). Our analysis showed that variations in CO ₂ fluxes were strongly correlated with microbial biomass pools, soil C and nitrogen (N) pools, soil inorganic P fluxes, dissolved organic C fluxes, and fine root biomass. Together, our data suggest that shifts in leaf litter inputs in response to localized human disturbances and global environmental change could have rapid and important consequences for belowground C storage and fluxes in tropical rain forests, and highlight differences between tropical and temperate ecosystems, where belowground C cycling responses to changes in litterfall are generally slower and more subtle.     

Nitrogen deposition,vegetation burning and climate warming act independently on microbial community structure and enzyme activity associated with decomposing litter in low‐alpine heath

Low‐alpine heathlands are thought to be particularly sensitive to nitrogen (N) deposition, climate and land management change, yet little is known about how these factors regulate key belowground processes, like litter turnover, under field conditions. Here we use an in situ factorial field experiment to test the effects of increased atmospheric N deposition, climate manipulation and past vegetation burning, and their interactions, on litter decomposition and the activity and diversity of associated microorganisms. The use of litter from within (native) and outwith (standard) the experimental plots also enabled us to test whether decomposition and microbial functional diversity is driven primarily by soil conditions or litter chemistry. In general, extracellular enzyme activities of litter were driven by additions of simulated N deposition with phosphatase being the most responsive. We found that standard litter incubated in plots that had been burnt 8 years previously decomposed slower and lost less N and phosphorus than in unburnt plots. This material also had associated with it the greatest activity of glucosidase and the least diverse microbial community, as assessed by culture‐independent methods. Although all treatments significantly affected microbial diversity, burning explained most of the variability, indicating a close coupling between plant and microbial communities in these treatments. A striking feature of all the data relating to both standard and native litter was an almost complete lack of interactive effects between the treatments. The lack of interactions between the treatments indicates that each perturbation might affect different mechanisms in the decomposition process (including the composition of associated microbial communities) and nutrient cycling.     

土壤微生物群落对枯落物输入的响应

枯落物分解在陆地生态系统物质循环能量流动中起着关键性作用,明确枯落物输入对土壤微生物群落的影响有助于理解土壤微生物生物多样性和陆地生态系统功能的相互关系。本文采用整合分析方法,以中国为研究区域,以不添加枯落物为对照组,探究土壤微生物(真菌、细菌、放线菌)及微生物生物量碳、生物量氮对枯落物输入的响应。结果表明:与不添加枯落物相比,添加枯落物后土壤微生物生物量碳、生物量氮分别显著增加3.9%和4.4%;土壤真菌PLFA、细菌PLFA及总微生物PLFA分别增加4.0%、3.1%和2.4%。枯落物输入对土壤微生物的影响受到气候条件、年降水量、植被类型及土壤酸碱度等因素的显著影响;不同气候类型下,土壤微生物对枯落物输入的响应呈现出亚热带季风气候区>温带季风气候区>温带大陆气候区的趋势,以及随着年降水量的增加呈现出先升高后降低的趋势;不同植被类型下,土壤微生物对枯落物输入的响应呈现出阔叶林>草地≈混交林>针叶林的趋势。     

Season mediates herbivore effects on litter and soil microbial abundance and activity in a semi-arid woodland

Herbivores can directly impact ecosystem function by altering litter quality of an ecosystem or indirectly by shifting the composition of microbial communities that mediate nutrient processes. We examined the effects of tree susceptibility and resistance to herbivory on litter microarthropod and soil microbial communities to test the general hypothesis that herbivore driven changes in litter inputs and soil microclimate will feedback to the microbial community. Our study population consisted of individual piñon pine trees that were either susceptible or resistant to the stem-boring moth (Dioryctria albovittella) and susceptible piñon pine trees from which the moth herbivores have been manually removed since 1982. Moth herbivory increased piñon litter nitrogen concentrations (16%) and decreased canopy precipitation interception (28%), both potentially significant factors influencing litter and soil microbial communities. Our research resulted in three major findings: (1) In spite of an apparent increase in litter quality, herbivory did not change litter microarthropod abundance or species richness. (2) However, susceptibility to herbivores strongly influenced bulk soil microbial communities (i.e., 52% greater abundance beneath herbivore-resistant and herbivore-removal trees than susceptible trees) and alkaline phosphatase activity (i.e., 412% increase beneath susceptible trees relative to other groups). (3) Season had a strong influence on microbial communities (i.e., microbial biomass and alkaline phosphatase activity increased after the summer rains), and their response to herbivore inputs, in this semi-arid ecosystem. Thus, during the dry season plant resistance and susceptibility to a common insect herbivore had little or no observable effects on the belowground organisms and processes we studied, but after the rains, some pronounced effects emerged.     

基于BIOLOG技术分析氮沉降和降水对土壤微生物功能多样性的影响

土壤微生物是有机物分解和养分循环的主要介质,因此在维持土壤的功能多样性和持续性方面发挥着关键作用。气候变化驱动因素会影响土壤微生物的生理活动,引起其群落结构和功能多样性的改变,并对生物地球化学循环和气候―生态系统反馈产生连锁效应,其中氮沉降和降水是全球气候变化的研究热点。土壤氮(N)的有效性有可能通过改变微生物的群落组成以调节微生物对降水变化的响应,但目前关于N沉降和降水及其交互作用对土壤微生物群落功能多样性的影响机制仍不清楚。为了准确预测未来气候条件下生态系统的功能状况,需要更好地了解土壤微生物对环境变化的响应。基于BIOLOG技术综述了氮沉降和降水变化及其交互作用对土壤微生物功能多样性影响的相关研究进展,可以为进一步研究全球气候变化背景下地下生态学的发展提供参考。另外,分析阐述了当前工作中存在的一些主要瓶颈,并对未来的研究热点进行了探讨和展望。     

Grass invasion causes rapid increases in ecosystem carbon and nitrogen storage in a semiarid shrubland

Accurately predicting terrestrial carbon (C) and nitrogen (N) storage requires understanding how plant invasions alter cycling and storage. A common, highly successful type of plant invasion occurs when the invasive species is of a distinctly different functional type than the native dominant plant, such as shrub encroachment throughout the western United States and annual grass invasions in Mediterranean shrublands, as studied here. Such invasions can dramatically transform landscapes and have large potential to alter C and N cycling by influencing storage in multiple pools. We used a manipulation of non‐native annual grass litter within a shrub‐dominated habitat in southern California (coastal sage scrub, CSS) to study how grass invasion alters ecosystem C and N storage. We added, removed, or left unchanged grass litter in areas of high and low invasion, then followed soil and vegetation changes. Grass litter greatly increased C and N storage in soil, aboveground native and non‐native biomass. Aboveground litter storage increased due to the greater inputs and slower decomposition of grass litter relative to shrub litter; shading by grass litter further reduced decomposition of both non‐native and native litter, which may be due to reduced photodegradation. Soil C and N pools in areas of high litter increased ～20% relative to low litter areas in the two years following manipulation and were generally sinks for C and N, while areas with low litter were sources. We synthesize our results into a C cycle of invaded and uninvaded areas of CSS and link changes in storage to increases in the soil fungi : bacteria ratio, increased plant inputs, and decreased litter loss. Overall, we show that grasses, especially through their litter, control important abiotic and biotic mechanisms governing C and N storage, with widespread implications for C sequestration and N storage in semiarid systems undergoing grass or shrub invasions.     

Ecosystem recovery: heathland response to a reduction in nitrogen deposition

Atmospheric deposition of nitrogen is responsible for widespread changes in the structure and function of sensitive seminatural ecosystems. The proposed reduction in emissions of nitrogenous pollutants in Europe under the Gothenburg Protocol raises the question of whether affected ecosystems have the potential to recover to their previous condition and, if so, over what timescale. Since 1998, we have monitored the response of a lowland heathland in southern England following the cessation of a long‐term nitrogen addition experiment, and subsequent management, assessing changes in vegetation growth and chemistry, soil chemistry and the soil microbial community. Persistent effects of earlier nutrient loading on Calluna growth and phenology, and on the abundance of lichens, were apparent up to 8 years after nitrogen additions ceased, indicating the potential for long‐term effects of modest nutrient loading (up to 15.4 kg N ha^?1 yr^?1, over 7 years) on heathland ecosystems. The size and activity of the soil microbial community was elevated in former N‐treated plots, 6–8 years after additions ceased, suggesting a prolonged effect on the rate of nutrient cycling. Although habitat management in 1998 reduced nitrogen stores in plant biomass, effects on belowground nitrogen stores were small. Although some parameters (e.g. soil pH) recover pretreatment levels relatively rapidly, others (e.g. vegetation cover and microbial activity) respond much more slowly, indicating that the ecological effects of even small increases in nitrogen deposition will persist for many years after deposition inputs are reduced. Indeed, calculations suggest that the additional soil nitrogen storage associated with 7 years of experimental nitrogen inputs could sustain the observed effects on plant growth and phenology for several decades. Carry over effects on plant phenology and sensitivity to drought suggest that the persistence of vegetation responses to nitrogen deposition should be integrated into long‐term assessments of the impact of global climate change on sensitive ecosystems.     

Linking Above- and Belowground Responses to 16 Years of Fertilization,Mowing, and Removal of the Dominant Species in a Temperate Grassland

Species-rich oligotrophic meadows are affected by a wide range of management interventions that influence their functioning and capacity to deliver ecosystem services, but long-term studies on the above- and belowground adaptations to different management tools are still scarce. We focused on the interactive effects of NPK fertilization, mowing, and removal of the initially dominant species (Molinia caerulea) on plant, soil, and microbial responses in wet oligotrophic grassland in a 16-year full-factorial manipulative experiment. Changes in vegetation composition, soil pH, and nutrient availability were accompanied by altered microbial phospholipid fatty acid (PLFA) composition, whereas treatment effects on soil microbial biomass and carbon (C) mineralization were mainly related to changes in soil organic matter (SOM) content and nutrient availability. Fertilization decreased plant species richness aboveground and lowered SOM storage and microbial activity belowground. Mowing preserved high plant diversity and led to more efficient recycling of N within the grassland, whereas Molinia removal significantly affected only plant community composition. Mowing combined with fertilization maintained high species richness only in the short term. Belowground, mowing reduced N leaching from the fertilized system but did not prevent SOM depletion, soil acidification, and concomitant adverse effects on soil microbes. We conclude that annual mowing is the appropriate type of extensive management for oligotrophic species-rich meadows, but the concomitant nutrient depletion should not be compensated for by regular NPK fertilization due to its adverse effects on soil quality.     

Interactive effects of vegetation type and elevation on aboveground and belowground properties in a subarctic tundra

An improved knowledge of how contrasting types of plant communities and their associated soil biota differ in their responses to climatic variables is important for better understanding the future impacts of climate change on terrestrial ecosystems. Elevational gradients serve as powerful study systems for answering questions on how ecological processes can be affected by changes in temperature and associated climatic variables. In this study, we evaluated how plant and soil microbial communities, and abiotic soil properties, change with increasing elevation in subarctic tundra in northern Sweden, for each of two dominant but highly contrasting vegetation types, namely heath (dominated by woody dwarf shrubs) and meadow (dominated by herbaceous species). To achieve this, we measured plant community characteristics, microbial community properties and several soil abiotic properties for both vegetation types across an elevation gradient of 500 to 1000 m. We found that the two vegetation types differed not only in several above‐ and belowground properties, but also in how these properties responded to elevation, pointing to important interactive effects between vegetation type and elevation. Specifically, for the heath, available soil nitrogen and phosphorus decreased with elevation whereas fungal dominance increased, while for the meadow, idiosyncratic responses to elevation for these variables were found. These differences in belowground responses to elevation among vegetation types were linked to shifts in the species and functional group composition of the vegetation. Our results highlight that these two dominant vegetation types in subarctic tundra differ greatly not only in fundamental aboveground and belowground properties, but also in how these properties respond to elevation and are therefore likely to be influenced by temperature. As such they highlight that vegetation type, and the soil abiotic properties that determine this, may serve as powerful determinants of how both aboveground and belowground properties respond to strong environmental gradients.     

菌根真菌影响森林生态系统碳循环研究进展

森林生态系统在全球碳(C)储量中占据极为重要的地位。菌根真菌广泛存在于森林生态系统中,在森林生态系统C循环过程中发挥重要的作用。阐述了不同菌根类型真菌在森林生态系统C循环过程中的功能,对比了温带/北方森林与热带/亚热带森林中菌根真菌介导的C循环研究方面新近取得的研究结果。发现温带和北方森林的外生菌根(EcM)植物对地上生物量C的贡献相对较小,然而是地下C储量的主要贡献者;以丛枝菌根(AM)共生为主的热带/亚热带森林地表生物量占比较高,表明AM植被对热带/亚热带森林地上生物量C的贡献相对较大。我们还就全球变化背景下,菌根真菌及其介导的森林生态系统C汇功能,以及不同菌根类型树种影响C循环的机制等进行了总结。菌根真菌通过影响凋落物分解、土壤有机质形成及地下根系生物量,进而影响整个森林生态系统的C循环功能。菌根介导的森林C循环过程很大程度上取决于(优势)树木的菌根类型和森林土壤中菌根真菌的群落结构。最后指出了当前研究存在的主要问题以及未来研究展望。本文旨在明确菌根真菌在森林生态系统C循环转化过程中的重要生态功能,有助于准确地评估森林生态系统C汇现状,为应对全球变化等提供重要的依据。     

Limited effects of early snowmelt on plants,decomposers, and soil nutrients in Arctic tundra soils

In addition to warming temperatures, Arctic ecosystems are responding to climate change with earlier snowmelt and soil thaw. Earlier snowmelt has been examined infrequently in field experiments, and we lack a comprehensive look at belowground responses of the soil biogeochemical system that includes plant roots, decomposers, and soil nutrients. We experimentally advanced the timing of snowmelt in factorial combination with an open‐top chamber warming treatment over a 3‐year period and evaluated the responses of decomposers and nutrient cycling processes. We tested two alternative hypotheses: (a) Early snowmelt and warming advance the timing of root growth and nutrient uptake, altering the timing of microbial and invertebrate activity and key nutrient cycling events; and (b) loss of insulating snow cover damages plants, leading to reductions in root growth and altered biological activity. During the 3 years of our study (2010–2012), we advanced snowmelt by 4, 15, and 10 days, respectively. Despite advancing aboveground plant phenology, particularly in the year with the warmest early‐season temperatures (2012), belowground effects were primarily seen only on the first sampling date of the season or restricted to particular years or soil type. Overall, consistent and substantial responses to early snowmelt were not observed, counter to both of our hypotheses. The data on soil physical conditions, as well interannual comparisons of our results, suggest that this limited response was because of the earlier date of snowmelt that did not coincide with substantially warmer air and soil temperatures as they might in response to a natural climate event. We conclude that the interaction of snowmelt timing with soil temperatures is important to how the ecosystem will respond, but that 1‐ to 2‐week changes in timing of snowmelt alone are not enough to drive season‐long changes in soil microbial and nutrient cycling processes.     

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

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

Root controls on soil microbial community structure in forest soils

We assessed microbial community composition as a function of altered above- and belowground inputs to soil in forest ecosystems of Oregon, Pennsylvania, and Hungary as part of a larger Detritus Input and Removal Treatment (DIRT) experiment. DIRT plots, which include root trenching, aboveground litter exclusion, and doubling of litter inputs, have been established in forested ecosystems in the US and Europe that vary with respect to dominant tree species, soil C content, N deposition rate, and soil type. This study used phospholipid fatty-acid (PLFA) analysis to examine changes in the soil microbial community size and composition in the mineral soil (0–10 cm) as a result of the DIRT treatments. At all sites, the PLFA profiles from the plots without roots were significantly different from all other treatments. PLFA analysis showed that the rootless plots generally contained larger quantities of actinomycete biomarkers and lower amounts of fungal biomarkers. At one of the sites in an old-growth coniferous forest, seasonal changes in PLFA profiles were also examined. Seasonal differences in soil microbial community composition were greater than treatment differences. Throughout the year, treatments without roots continued to have a different microbial community composition than the treatments with roots, although the specific PLFA biomarkers responsible for these differences varied by season. These data provide direct evidence that root C inputs exert a large control on microbial community composition in the three forested ecosystems studied.     

Are patterns in nutrient limitation belowground consistent with those aboveground: results from a 4 million year chronosequence

Accurately predicting the effects of global change on net carbon (C) exchange between terrestrial ecosystems and the atmosphere requires a more complete understanding of how nutrient availability regulates both plant growth and heterotrophic soil respiration. Models of soil development suggest that the nature of nutrient limitation changes over the course of ecosystem development, transitioning from nitrogen (N) limitation in ‘young’ sites to phosphorus (P) limitation in ‘old’ sites. However, previous research has focused primarily on plant responses to added nutrients, and the applicability of nutrient limitation-soil development models to belowground processes has not been thoroughly investigated. Here, we assessed the effects of nutrients on soil C cycling in three different forests that occupy a 4 million year substrate age chronosequence where tree growth is N limited at the youngest site, co-limited by N and P at the intermediate-aged site, and P limited at the oldest site. Our goal was to use short-term laboratory soil C manipulations (using ¹⁴C-labeled substrates) and longer-term intact soil core incubations to compare belowground responses to fertilization with aboveground patterns. When nutrients were applied with labile C (sucrose), patterns of microbial nutrient limitation were similar to plant patterns: microbial activity was limited more by N than by P in the young site, and P was more limiting than N in the old site. However, in the absence of C additions, increased respiration of native soil organic matter only occurred with simultaneous additions of N and P. Taken together, these data suggest that altered nutrient inputs into ecosystems could have dissimilar effects on C cycling above- and belowground, that nutrients may differentially affect of the fate of different soil C pools, and that future changes to the net C balance of terrestrial ecosystems will be partially regulated by soil nutrient status.     

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.     

Impacts of altered precipitation regimes on soil communities and biogeochemistry in arid and semi‐arid ecosystems

Altered precipitation patterns resulting from climate change will have particularly significant consequences in water‐limited ecosystems, such as arid to semi‐arid ecosystems, where discontinuous inputs of water control biological processes. Given that these ecosystems cover more than a third of Earth's terrestrial surface, it is important to understand how they respond to such alterations. Altered water availability may impact both aboveground and belowground communities and the interactions between these, with potential impacts on ecosystem functioning; however, most studies to date have focused exclusively on vegetation responses to altered precipitation regimes. To synthesize our understanding of potential climate change impacts on dryland ecosystems, we present here a review of current literature that reports the effects of precipitation events and altered precipitation regimes on belowground biota and biogeochemical cycling. Increased precipitation generally increases microbial biomass and fungal:bacterial ratio. Few studies report responses to reduced precipitation but the effects likely counter those of increased precipitation. Altered precipitation regimes have also been found to alter microbial community composition but broader generalizations are difficult to make. Changes in event size and frequency influences invertebrate activity and density with cascading impacts on the soil food web, which will likely impact carbon and nutrient pools. The long‐term implications for biogeochemical cycling are inconclusive but several studies suggest that increased aridity may cause decoupling of carbon and nutrient cycling. We propose a new conceptual framework that incorporates hierarchical biotic responses to individual precipitation events more explicitly, including moderation of microbial activity and biomass by invertebrate grazing, and use this framework to make some predictions on impacts of altered precipitation regimes in terms of event size and frequency as well as mean annual precipitation. While our understanding of dryland ecosystems is improving, there is still a great need for longer term in situ manipulations of precipitation regime to test our model.     

Legacy effects of aboveground-belowground interactions

Root herbivory can greatly affect the performance of aboveground insects via changes in plant chemistry. These interactions have been studied extensively in experiments where aboveground and belowground insects were feeding on the same plant. However, little is known about how aboveground and belowground organisms interact when they feed on plant individuals that grow after each other in the same soil. We show that feeding by aboveground and belowground insect herbivores on ragwort (Jacobaea vulgaris) plants exert unique soil legacy effects, via herbivore-induced changes in the composition of soil fungi. These changes in the soil biota induced by aboveground and belowground herbivores of preceding plants greatly influenced the pyrrolizidine alkaloid content, biomass and aboveground multitrophic interactions of succeeding plants. We conclude that plant-mediated interactions between aboveground and belowground insects are also important when they do not feed simultaneously on the same plant.