Belowground Ecosystem Research Articles

Linking soil food web structure to above- and belowground ecosystem processes: a meta-analysis

Soil fauna can be an important regulator of community parameters and ecosystem processes, but there have been few quantitative syntheses of the role of soil fauna in terrestrial soil communities and ecosystems. Here, we conducted a meta-analysis to investigate the impacts of invertebrate soil micro- and mesofauna (grazers and predators) on plant productivity and microbial biomass. Overall our results indicate that an increase in the biomass of soil fauna increased aboveground plant productivity across ecosystems by 35% and decreased microbial biomass by 8%. In addition, we found no evidence for trophic cascades in terrestrial soil food webs, but the bacterivorous component of soil fauna influenced plant productivity and microbial biomass more than did the fungivorous component. Furthermore, changes in the biomass of soil fauna differentially affected plant productivity among plant functional groups: a higher biomass of soil fauna increased aboveground productivity by 70% in coniferous systems. However, in ecosystems dominated by legumes, a functional group with lower inorganic nitrogen requirements, there was no response of aboveground productivity to increases in the biomass of soil fauna. In sum, the results of this meta-analysis indicate that soil fauna help to regulate ecosystem production, especially in nutrient-limited ecosystems.

Biomass, and carbon and nitrogen pools in a subtropical evergreen broad-leaved forest in eastern China

Subtropical evergreen broad-leaved forest is the most widely distributed land-cover type in eastern China. As the rate of land-use change accelerates worldwide, it is becoming increasingly important to quantify ecosystem biomass and carbon (C) and nitrogen (N) pools. Above and below-ground biomass and ecosystem pools of N and C in a subtropical secondary forest were investigated at Laoshan Mountain Natural Reserve, eastern China. Total biomass was 142.9 Mg ha−1 for a young stand (18 years) and 421.9 Mg ha−1 for a premature stand (ca. 60 years); of this, root biomass was from 26.9 (18.8% of the total) to 100.3 Mg ha−1 (23.8%). Total biomass C and N pools were, respectively, 71.4 Mg ha−1 and 641.6 kg ha−1 in the young stand, and 217.0 Mg ha−1 and 1387.4 kg ha−1 in the premature stand. The tree layer comprised 91.8 and 89.4% of the total biomass C and N pools in the young stand, and 98.0 and 95.6% in the premature stand. Total ecosystem C and N pools were, respectively, 101.4 and 4.6 Mg ha−1 for the young stand, and 260.2 and 6.6 Mg ha−1 for the premature stand. Soil C comprised 23.8–29.6% of total ecosystem C whereas soil N comprised 76.9–84.4% of the total. Our results suggest that a very high percentage of N in this subtropical forest ecosystem is stored in the mineral soil, whereas the proportion of organic C in the soil pool is more variable. The subtropical forest in eastern China seems to rapidly accumulate biomass during secondary succession, which makes it a potentially rapid accumulator of, and large sink for, atmospheric C.

A review on the eco-geography of soil fauna in China

This review summarizes main research findings in soil fauna eco-geography in China in the past 30 years. The subject areas and main results were overviewed including biodiversity and eco-geological distribution of soil fauna communities. Studies of ecological distributions of soil fauna and dynamic ranges in space from tropical, subtropical to temperate regions, and in categories from forest, grassland, desert, wetland, farmland to urban ecosystems, the responses and indications of soil fauna to soil environments. Effects of intensive disturbance such as fire, grazing, farming, fertilization on soil fauna include sensitive (e.g. nematode) and rare groups and community indexes. The functions of soil fauna were discussed including environmental construction, environmental purification, litter decomposition and elements cycling. Interactions between soil fauna and other biota in soil ecosystems and linking between aboveground and belowground diversity and the effects of global change on soil fauna community in China were also included. Finally, the authors pointed out common interests in soil fauna eco-geographical studies, which include application of molecule biology into soil fauna taxa; function and mechanism of soil fauna community diversity; interaction between aboveground and belowground ecosystems; effects of disturbance, pollution, biological invasion, and global change on soil fauna community and function. The review is to provide a scientific basis for promoting soil fauna eco-geographical studies in China.

Responses of soil chemical and biological properties to nitrogen addition in a Dahurian larch plantation in Northeast China

Soil microbial properties play a key role in belowground ecosystem functioning, but are not well understood in forest ecosystems under nitrogen (N) enrichment. In this study, soil samples from 0–10 cm and 10–20 cm layers were collected from a Dahurian larch (Larix gmelinii Rupr.) plantation in Northeast China after six consecutive years of N addition to examine changes in soil pH, nutrient concentrations, and microbial biomass and activities. Nitrogen addition significantly decreased soil pH and total phosphorus, but had little effect on soil total organic carbon (TOC) and total N (TN) concentrations. The NO3−-N concentrations in the two soil layers under N addition were significantly higher than that in the control, while NH4+-N concentrations were not different. After six years of N addition, potential net N mineralization and nitrification rates were dramatically increased. Nitrogen addition decreased microbial biomass C (MBC) and N (MBN), and MBC/TOC and MBN/TN in the 0–10 cm soil layer, but MBC/MBN was increased by 67% in the 0–10 cm soil layer. Soil basal respiration, microbial metabolic quotient (qCO2), and β-glucosidase, urease, acid phosphomonoesterase and nitrate reductase activities in the two soil layers showed little change after six years of N addition. However, soil protease and dehydrogenase activities in the 0–10 cm layer were 41% and 54% lower in the N addition treatment than in the control, respectively. Collectively, our results suggest that in the mid-term N addition leads to a decline in soil quality in larch plantations, and that different soil enzymes show differentiated responses to N addition.

Effects of biological invasions on forest carbon sequestration

AbstractThere has been a rapidly developing literature on the effects of some of the major drivers of global change on carbon (C) sequestration, particularly carbon dioxide (CO2) enrichment, land use change, nitrogen (N) deposition and climate change. However, remarkably little attention has been given to one major global change driver, namely biological invasions. This is despite growing evidence that invasive species can dramatically alter a range of aboveground and belowground ecosystem processes, including those that affect C sequestration. In this review, we assess the evidence for the impacts of biological invaders on forest C stocks and C sequestration by biological invaders. We first present case studies that highlight a range of invader impacts on C sequestration in forest ecosystems, and draw on examples that involve invasive primary producers, decomposers, herbivores, plant pathogens, mutualists and predators. We then develop a conceptual framework for assessing the effects of invasive species on C sequestration impacts more generally, by identifying the features of biological invaders and invaded ecosystems that are thought to most strongly regulate C in forests. Finally we assess the implications of managing invasive species on C sequestration. An important principle that emerges from this review is that the direct effects of invaders on forest C are often smaller and shorter‐term than their indirect effects caused by altered nutrient availability, primary productivity or species composition, all of which regulate long‐term C pools and fluxes. This review provides a conceptual basis for improving our general understanding of biological invaders on ecosystem C, but also points to a paucity of primary data that are needed to determine the quantitative effects of invaders on ecosystem processes that drive C sequestration.

Plant-soil feedback of native and range-expanding plant species is insensitive to temperature.

Temperature change affects many aboveground and belowground ecosystem processes. Here we investigate the effect of a 5°C temperature increase on plant–soil feedback. We compare plant species from a temperate climate region with immigrant plants that originate from warmer regions and have recently shifted their range polewards. We tested whether the magnitude of plant–soil feedback is affected by ambient temperature and whether the effect of temperature differs between these groups of plant species. Six European/Eurasian plant species that recently colonized the Netherlands (non-natives), and six related species (natives) from the Netherlands were selected. Plant–soil feedback of these species was determined by comparing performance in conspecific and heterospecific soils. In order to test the effect of temperature on these plant–soil feedback interactions, the experiments were performed at two greenhouse temperatures of 20/15°C and 25/20°C, respectively. Inoculation with unconditioned soil had the same effect on natives and non-natives. However, the effect of conspecific conditioned soil was negative compared to heterospecific soil for natives, but was positive for non-natives. In both cases, plant–soil interactions were not affected by temperature. Therefore, we conclude that the temperature component of climate change does not affect the direction, or strength of plant–soil feedback, neither for native nor for non-native plant species. However, as the non-natives have a more positive soil feedback than natives, climate warming may introduce new plant species in temperate regions that have less soil-borne control of abundance.

Missing links in the root–soil organic matter continuum

The soil environment remains one of the most complex and poorly understood research frontiers in ecology. Soil organic matter (SOM), which spans a continuum from fresh detritus to highly processed, mineral-associated organic matter, is the foundation of sustainable terrestrial ecosystems. Heterogeneous SOM pools are fueled by inputs from living and dead plants, driven by the activity of micro- and mesofauna, and are shaped by a multitude of abiotic factors. The specialization required to measure unseen processes that occur on a wide range of spatial and temporal scales has led to the partitioning of soil ecology research across several disciplines. In the organized oral session 'Missing links in the root-soil organic matter continuum' at the annual Ecological Society of America meeting in Albuquerque, NM, USA, we joined the call for greater communication and collaboration among ecologists who work at the root-soil interface (e.g. Coleman, 2008). Our goal was to bridge the gap between scientific disciplines and to synthesize disconnected pieces of knowledge from root-centric and soil-centric studies into an integrated understanding of belowground ecosystem processes. We focused this report around three compelling themes that arose from the session: (1) the influence of the rhizosphere on SOM cycling, (2) the role ofmore » soil heterotrophs in driving the transformation of root detritus to SOM, and (3) the controlling influence of the soil environment on SOM dynamics. We conclude with a discussion of new approaches for gathering data to bridge gaps in the root-SOM continuum and to inform the next generation of ecosystem models. Although leaf litter has often been considered to be the main source of organic inputs to soil, Ann Russell synthesized a convincing body of work demonstrating that roots, rather than surface residues, control the accumulation of SOM in a variety of ecosystems. Living roots, which are chemically diverse and highly dynamic, also influence a wide range of soil processes, from the exudation of labile C compounds to the development of fungal associations. For example, Zoe Cardon demonstrated that the root-mediated redistribution of deep soil water to relatively dry shallower soil, increased soil CO{sub 2} efflux and nutrient cycling near the surface in an arid ecosystem. Andrew Kulmatiski also discussed the importance of rooting distribution throughout the soil profile for strategies of water uptake by different species in an African savanna. Later, Julie Jastrow demonstrated that living roots shape soil physical structure by promoting the formation of soil aggregates, which facilitated accrual of SOM in restored grasslands. Taken together, the evidence is compelling that living roots, and organic matter derived from root detritus, are important parts of the continuum of organic matter in the soil. Larger soil organisms (i.e. 50 {micro}m to many cm in body size) play an important role in the root-SOM continuum by grazing on roots and microbes, comminuting organic matter and aggregating soil in fecal pellets. However, litterbag and soil incubation studies necessarily exclude invertebrates, and research on faunal activity and trophic dynamics tends to be independent from research on the biogeochemistry of SOM cycling. Tim Filley used plant-derived biomarkers in invertebrate residues to bridge the gap between larger soil organisms, such as earthworms and beetle larvae, and SOM distribution. He found that larger soil organisms help to stabilize root-derived organic matter in soil aggregates. Similar coupling of biogeochemistry with food web studies could prove fruitful for describing mechanisms that underlie critical ecosystem processes. Despite considerable research efforts, the breadth of the microbial role in the root-SOM continuum remains unresolved. Using advanced pyrosequencing techniques, David Nelson demonstrated the importance of archea as nitrifiers in agricultural systems exposed to elevated [CO{sub 2}]. Rising atmospheric [CO{sub 2}] and other changing environmental factors add a layer of complexity to the quest to understand microbial process. For example, Claudia Boot demonstrated that microbially mediated C and N cycling in Mediterranean California grasslands is intricately linked with summer drought. Ongoing research across subdisciplines seeks to uncover the many complex links between soil organisms of all sizes and the root-SOM continuum. While the role that living organisms play in the transformation of root detritus to SOM is disproportionate to their body size, the nonliving soil environment also influences SOM cycling. However, destructive sampling can obscure feedbacks between abiotic and biotic processes, making it difficult to quantify the role of edaphic factors in the root-SOM continuum.« less

Global patterns in belowground communities

Although belowground ecosystems have been studied extensively and soil biota play integral roles in biogeochemical processes, surprisingly we have a limited understanding of global patterns in belowground biomass and community structure. To address this critical gap, we conducted a meta-analysis of published data (> 1300 datapoints) to compare belowground plant, microbial and faunal biomass across seven of the major biomes on Earth. We also assembled data to assess biome-level patterns in belowground microbial community composition. Our analysis suggests that variation in microbial biomass is predictable across biomes, with microbial biomass carbon representing 0.6-1.1% of soil organic carbon (r(2) = 0.91) and 1-20% of total plant biomass carbon (r(2) = 0.42). Approximately 50% of total animal biomass can be found belowground and soil faunal biomass represents < 4% of microbial biomass across all biomes. The structure of belowground microbial communities is also predictable: bacterial community composition and fungal : bacterial gene ratios can be predicted reasonably well from soil pH and soil C : N ratios respectively. Together these results identify robust patterns in the structure of belowground microbial and faunal communities at broad scales which may be explained by universal mechanisms that regulate belowground biota across biomes.

Exotic Ecosystem Engineers Change the Emergence of Plants from the Seed Bank of a Deciduous Forest

The anthropogenic spread of exotic ecosystem engineers profoundly impacts native ecosystems. Exotic earthworms were shown to alter plant community composition of the understory of deciduous forests previously devoid of earthworms. We investigated the effect of two exotic earthworm species (Lumbricus terrestris L. and Octolasion tyrtaeum Savigny) belonging to different ecological groups (anecic and endogeic) on the emergence of plants from the seed bank of a northern North American deciduous forest using the seedling emergence method. We hypothesized that (1) exotic earthworms change the seedling emergence from the plant seed bank, (2) L. terrestris increases the emergence of plant seedlings of the deeper soil layer but decreases that of the upper soil layer due to plant seed burial, and (3) O. tyrtaeum decreases plant seedling emergence due the damage of plant seeds. Indeed, exotic earthworms altered the emergence of plant seedlings from the seed bank and the functional composition of the established plant seedlings. Surprisingly, although L. terrestris only marginally affected seedling emergence, O. tyrtaeum changed the emergence of native plant species from the seed bank considerably. In particular, the number of emerging grass and herb seedlings were increased in the presence of O. tyrtaeum in both soil layers. Moreover, the impacts of earthworms depended on the identity of plant functional groups; herb species benefited, whereas legumes suffered from the presence of exotic earthworms. The results highlight the strong effect of invasive belowground ecosystem engineers on aboveground ecosystem characteristics and suggest fundamental changes of ecosystems by human-spread earthworm species.

Carbon fluxes and nitrogen availability along an urban–rural gradient in a desert landscape

Urbanization is increasing in arid and semi-arid regions of the world and impacting native ecosystems through disturbance, climatic modification, and pollution deposition. Arid ecosystems often exhibit a mosaic of shrub/tree canopy covered and non-canopy covered patches that differ in elemental pools and processes. We measured belowground ecosystem attributes and processes in native Larrea tridentata {(D.C.) Cov} dominated communities along an urban–rural gradient in Phoenix, Arizona. Organic carbon (C), total nitrogen (N), and nitrate levels were significantly greater in the urban locations, but soil respiration rates (Rs) were higher at the rural sites. Urban sites exhibited no difference in Rs and N between the canopy and interplant patches while both the rural and suburban sites had significantly greater N and higher Rs under the canopy than in the interplant spaces. Soil respiration rates at the urban locations were not correlated with either soil moisture or temperature individually. These data suggest that urbanization has caused a disconnect of ecological pattern and processes in L. tridentata ecosystems within the urban setting such that water and N limitations no longer explain expected spatial Rs patterns, or elemental pools.

Hydraulic redistribution may stimulate decomposition

Roots influence root litter decomposition through multiple belowground processes. Hydraulic lift or redistribution (HR) by plants is one such process that creates diel drying–rewetting cycles in soil. However, it is unclear if this phenomenon influences decomposition. Since decomposition in deserts is constrained by low soil moisture and is stimulated when dry soils are rewetted, we hypothesized that diel drying–rewetting, via HR, stimulates decomposition of root litter. We quantified the decomposition of root litter from two desert shrubs, Artemisia tridentata ssp. tridentata and Sarcobatus vermiculatus, during spring and summer in field soil core treatments designed to have abundant roots and high magnitude HR cycles (DenseRoot) or few roots and low magnitude HR (SparseRoot). To help explain our decomposition results, we not only evaluated HR, but multiple factors (i.e., soil moisture, soil temperature, dissolved soil organic C concentrations, and litter chemistry) that are often influenced by roots and regulate decomposition. Root length density in the DenseRoot treatment was at least four times higher than in the SparseRoot treatment for both Artemisia and Sarcobatus by the beginning of spring. During spring and summer, there was only one instance when decomposition rates differed between the treatments. This occurred in soils beneath Artemisia in the summer when decomposition rates were 25% higher in the DenseRoot than in the SparseRoot treatments. Of the factors evaluated, only a threefold increase in the magnitude of drying–rewetting cycles created by HR in the DenseRoot compared to the SparseRoot treatment coincided with this change in decomposition. Additionally, the lower soil Ψw present in the Artemisia DenseRoot treatment should have resulted in a decline in decomposition rates, but the presence of higher magnitude HR cycles seemed to nullify this effect. There was no evidence of this result in Sarcobatus soils, possibly due to Sarcobatus only creating HR cycles for a short period of time in the summer before soil Ψw dropped below −7 MPa. As hypothesized, our results suggest that the presence of high magnitude HR cycles stimulated decomposition. The most plausible mechanism for this stimulation; however, was not solely due to HR drying–rewetting cycles but HR creating a diel rhythm of root-driven water fluxes and rhizodeposition. These together heightened microbial activity and, subsequently, enhanced the decomposition of surrounding litter. Our findings are the first field data supporting suggestions that HR influences belowground ecosystem processes and demonstrates that this relationship is seasonally variable.

Effect of ecosystem retrogression on stable nitrogen and carbon isotopes of plants, soils and consumer organisms in boreal forest islands

In the prolonged absence of catastrophic disturbance, ecosystem retrogression occurs, and this involves increased nutrient limitation, and reduced aboveground and belowground ecosystem processes rates. Little is known about how the nitrogen and carbon stable isotope ratios (delta(15)N and delta(13)C) of plants, soils and consumer organisms respond to retrogression in boreal forests. We investigated a 5000 year chronosequence of forested islands in the boreal zone of northern Sweden, for which the time since lightning-induced wildfire increases with decreasing island size, leading to ecosystem retrogression. For this system, tissue delta(15)N of three abundant plant species (Betula pubescens, Vaccinium myrtillus and Pleurozium schreberi) and humus all increased as retrogression proceeded. This is probably due to enhanced ecosystem inputs of N by biological fixation, and greater dependency of the plants on organic N during retrogression. The delta(13)C of B. pubescens and plant-derived humus also increased during retrogression, probably through nutrient limitation increasing plant physiological stress. Unlike the plants, delta(15)N of invertebrates (lycosid spiders and ants) did not increase during retrogression, probably because of their partial dependence on aquatic-derived prey that had a variable delta(15)N signature. The delta(13)C of the invertebrates increased as retrogression proceeded and converged towards that of an aquatic prey source (chironomid flies), suggesting increased dependence on aquatic-derived prey during retrogression. These results show that measurement of delta(15)N and delta(13)C of plants, soils, and consumers across the same environmental gradient can provide insights into environmental factors that drive both the aboveground and belowground subsystems, as well as the linkages between them.

Flooding effects on soil microbial communities

Flooding affects both above- and below-ground ecosystem processes. While the below-ground changes may be less obvious, they are as important as the above-ground changes. Soil microorganisms are sensitive to disturbance, and shifts in soil microbial community structure are expected when anaerobic conditions develop from flooding. The primary objective of these studies was to determine the effect of flooding on soil microbial communities. Simulated floods were established under greenhouse and field conditions. Flood treatments of flowing, intermittent (greenhouse only) or stagnant conditions were compared to a control with no flooding. In addition, residue treatments (incorporation of grass, legume or tree residue) were evaluated under greenhouse conditions. Soil samples from these experiments were examined using phospholipid fatty acid (PLFA) analysis and total N (TN), total organic carbon (TOC) and C:N ratio. Stagnant flood conditions in the greenhouse decreased microbial biomass and markers for aerobic bacteria, Gram-negative bacteria, Gram-positive bacteria, and mycorrhizal fungi. However, residue treatment, in general, did not affect microbial community structure. Effects of flood treatments in the field varied with depth and flood × depth interaction. The B:F ratio and microbial biomass decreased with stagnant flooding while other measures were not affected by flooding. Microbial biomass and microbial markers decreased with depth. We found some changes in the soil microbial community due to flooding; however, the 5-week time period of our study may not have been long enough to develop measureable changes. Further changes in the microbial community may occur as flood waters remain in a given area.

Links between plant litter chemistry, species diversity, and below-ground ecosystem function

Decomposition is a critical source of plant nutrients, and drives the largest flux of terrestrial C to the atmosphere. Decomposing soil organic matter typically contains litter from multiple plant species, yet we lack a mechanistic understanding of how species diversity influences decomposition processes. Here, we show that soil C and N cycling during decomposition are controlled by the composition and diversity of chemical compounds within plant litter mixtures, rather than by simple metrics of plant species diversity. We amended native soils with litter mixtures containing up to 4 alpine plant species, and we used 9 litter chemical traits to evaluate the chemical composition (i.e., the identity and quantity of compounds) and chemical diversity of the litter mixtures. The chemical composition of the litter mixtures was the strongest predictor of soil respiration, net N mineralization, and microbial biomass N. Soil respiration and net N mineralization rates were also significantly correlated with the chemical diversity of the litter mixtures. In contrast, soil C and N cycling rates were poorly correlated with plant species richness, and there was no relationship between species richness and the chemical diversity of the litter mixtures. These results indicate that the composition and diversity of chemical compounds in litter are potentially important functional traits affecting decomposition, and simple metrics like plant species richness may fail to capture variation in these traits. Litter chemical traits therefore provide a mechanistic link between organisms, species diversity, and key components of below-ground ecosystem function.

Towards a predictive understanding of belowground process responses to climate change: have we moved any closer?

Summary 1. Belowground processes, including root production and exudation, microbial activity and community dynamics, and biogeochemical cycling interact to help regulate climate change. Feedbacks associated with these processes, such as warming-enhanced decomposition rates, give rise to major uncertainties in predictions of future climate. 2. Uncertainties associated with these processes are more likely to be reduced if two key challenges can be met: increasing interdisciplinarity among researchers, and measuring belowground ecosystem structure and function at relevant spatial and temporal scales. For instance, recognizing the relationship between belowground primary production and soil respiration enhances modelling of global-scale C cycle temperature responses. At the opposite end of the spectrum, applying genomic techniques at the scale of microns improves mechanistic understanding of root‐microbe interactions. 3. Progress has been made in understanding interactions of belowground processes with climate change, although challenges remain. We highlight some of these advances and provide directions for key research needs in this Special Feature of Functional Ecology, which results from a symposium that was convened at the Soil Science Society of America National Meeting in November, 2006.

Context dependency of litter‐mixing effects on decomposition and nutrient release across a long‐term chronosequence

Litter decomposition is an important driver of terrestrial systems, and factors that determine decomposition rate for individual litter species have been widely studied. Fewer studies have explored the factors that regulate how mixing litters of multiple species affects litter decomposition and nutrient dynamics, and only a handful of studies have investigated how litter‐mixing effects may differ among different habitats or ecosystems, or how they respond to environmental gradients. We used a well‐established retrogressive chronosequence involving thirty lake islands in northern Sweden in which time since fire disturbance increases with decreasing island size; smaller islands therefore have reduced rates of aboveground and belowground ecosystem processes. On each of these islands we utilized plots with and without the long‐term experimental removal of shrubs. Litters from the six most common plant species on the islands were prepared in single‐, three‐ and six‐species litterbags, and placed on both the shrub‐removal and non‐removal plots on each island to decompose for one year. We found significant non‐additive effects of litter mixing on litter decomposition rates, on final litter N and P concentrations, and on litter N loss, but these non‐additive effects varied both in direction and magnitude with changed number of species, and even among litter mixtures with the same number of species. Further, the magnitude of non‐additive effects of litter mixing on both litter decomposition and nutrient dynamics was significantly influenced by both island size and the interaction between island size and shrub‐removal treatment. When shrubs were present, there was a U‐shaped relationship between these non‐additive effects and island size, while the relationship was positive when shrubs were removed. Hence, our results support previous findings that litter mixing may produce non‐additive effects on litter decomposition and nutrient dynamics, and that these effects tend to be idiosyncratic due to the importance of effects of individual species in the mixture. Most importantly, our results show that non‐additive litter‐mixing effects change greatly across environmental gradients, meaning that the biotic and abiotic characteristics of an ecosystem can be a powerful driver of the magnitude and even the direction of litter‐mixing effects on ecosystem processes.

Aggregate and organic matter dynamics in reclaimed soils as indicated by stable carbon isotopes

Recovery of belowground ecosystem processes, such as soil aggregation and organic matter (OM) accumulation, in reconstructed soils is crucial to successful reclamation of disturbed lands. Objectives of this study were to track soil aggregate recovery in combination with aggregate associated OM on a chronosequence of reclaimed surface mine sites and a native, undisturbed reference site. Macroaggregate and micro-within-macroaggregate proportions increased with reclamation age, while microaggregate proportions decreased. Organic carbon (C) and total nitrogen (N) concentrations increased with reclamation age for each aggregate fraction and were higher in the OM fraction observed within soil aggregates than in the free OM fraction found between soil aggregates. Naturally occurring isotopic signatures of 13C decreased rapidly with reclamation age, indicating over 50% of total aggregate C to be new C from predominately C 3 plant community inputs after 26 years of reclamation. Soil aggregate size distribution trends of increasing macroaggregation and micro-within-macroaggregates along with rapid rates of OM accumulation with time indicated that reclaimed soils had recovered structurally towards a native soil condition after a period of 10–15 years.

Invasion by an exotic tree alters above and belowground ecosystem components

With the widespread introduction and invasion of exotic plants there is a need for studies that quantify alterations of basic ecosystem structure and function. Ecosystem invasion by Melaleuca quinquenervia significantly altered both above- and belowground ecosystem components in this study. We measured the quantity and nutrient concentration of the litterfall, litter layer, and soil; microbial biomass pools; and rates of potentially mineralizable nitrogen and soil oxygen demand. Annual litterfall was 4.9 times higher in the non-invaded sites and contained 1.9 times more phosphorus than invaded sites. Non-invaded plots contained a larger litter layer compared to invaded plots: 2.4 ± 1.2 kg m−2 and 0.62 ± 0.3 kg m−2, respectively. Lower nutrient concentration and quantity of the litter layer in the invaded plots led to changes in the aboveground storage of nutrients. In the invaded plots there was four times less carbon, seven times less nitrogen, and ten times less phosphorus stored in the organic litter layer compared to the non-invaded plots. Microbial biomass nutrient pools were consistently lower at both the 0–5 cm and 5–15 cm depth in the invaded soils compared to non-invaded soils, indicating a plant mediated change. Although M. quinquenervia altered microbial community structure, microbial activities were not different between invaded and non-invaded plots at either depth as measured by rates of soil oxygen demand and potentially mineralizable nitrogen. These changes may affect both native plant growth and water quality, and may act to promote and maintain site dominance by M. quinquenervia.

Long term repeated burning in a wet sclerophyll forest reduces fungal and bacterial biomass and responses to carbon substrates

Soils from a long term experiment, established in 1972, incorporating replicated treatments of burning every 2 and 4 years with control plots were sampled in 2005 to determine the changes in microbial community structure, measured using phospholipid fatty acids (PLFAs) and functional diversity measured using multiple substrate induced respiration (SIR) tests (MicroResp™). Microbial biomass (total PLFA) in the 2 year burn treatments was 50% less than both the control and 4-year burn treatments. There was also concomitantly less respiratory activity which mirrored the known changes in soil C and substrate quality. Contrary to other studies soil bacterial PLFAs were reduced as much as fungal PLFAs in the 2-year burn and the short term (6 h) SIR of arginine, lysine, galactose and trehalose were significantly inhibited in the 2-year burn soils. The data suggest that a 4-year burn is a more sustainable practice for maintaining the original structure and function of the forest belowground ecosystem.

Shrub-interspace dynamics alter relationships between microbial community composition and belowground ecosystem characteristics

In desert ecosystems, belowground characteristics are influenced chiefly by the formation and persistence of “shrub-islands of fertility” in contrast to barren plant interspaces. If soil microbial communities are exclusively compared between these two biogeochemically distinct soil types, the impact of characteristics altered by shrub species, especially soil C and N, are likely to be overemphasized and overshadow the role of other characteristics in structuring microbial composition. To determine how belowground characteristics influence microbial community composition, and if the relative importance of these characteristics shifts across the landscape (i.e., between and within shrub and interspace soils), changes in microbial communities across a 3000-year cold desert chronosequence were related to 27 belowground characteristics in surface and subsurface soils. When shrub and interspace communities in surface and subsurface soils were combined across the entire chronosequence, communities differed and were primarily influenced by soil C, NO 3 − concentrations, bulk density, pH, and root presence. Within shrub soils, microbial communities were shrub species-specific, especially in surface soils, highlighting differences in soil characteristics created by specific shrub species and/or similarity in stresses structuring shrub species and microbial communities alike. Microbial communities in shrub soils were not influenced by soil C, but by NO 3 − and NH 4 + concentrations, pH, and silt in surface soils; and Cl, P, soil N, and NO 3 − concentrations in subsurface soils. Interspace soil communities were distinct across the chronosequence at both depths and were strongly influenced by dune development. Interspace communities were primarily associated with soil stresses (i.e., high B and Cl concentrations), which decreased with dune development. The distribution of Gram-positive bacteria, Actinobacteria, and fungi highlighted community differences between and within shrub and interspace soils, while Gram-negative bacteria were common in all soils across the chronosequence. Of the 27 belowground characteristics investigated, 13 separated shrub from interspace communities, and of those, only five emerged as factors influencing community composition within shrub and interspace soils. As dunes develop across this cold desert chronosequence, microbial community composition was not regulated primarily by soil C, but by N and P availability and soil stresses in shrub soils, and exclusively by soil stresses in interspace soils.