Impact of litter quality on mineralization processes in managed and abandoned pasture soils in Southern Ecuador

Tropical regions are currently undergoing remarkable rates of land use change accompanied by altered litter inputs to soil. In vast areas of Southern Ecuador forests are clear cut and converted for use as cattle pastures. Frequently these pasture sites are invaded by bracken fern, when bracken becomes dominant pasture productivity decreases and the sites are abandoned. In the present study implications of invasive bracken on soil biogeochemical properties were investigated. Soil samples (0-5 cm) were taken from an active pasture with Setaria sphacelata as predominant grass and from an abandoned pasture overgrown by bracken. Grass (C₄ plant) and bracken (C₃ plant) litter, differing in C:N ratio (33 and 77, respectively) and lignin content (Klason-lignin: 18% and 45%, respectively), were incubated in soils of their corresponding sites and vice versa for 28 days at 22 °C. Unamended microcosms containing only the respective soil or litter were taken as controls. During incubation the amount of CO₂ and its δ¹³C-signature were determined at different time intervals. Additionally, the soil microbial community structure (PLFA-analysis) as well as the concentrations of KCl-extractable C and N were monitored. The comparison between the control soils of active and abandoned pasture sites showed that the massive displacement of Setaria-grass by bracken after pasture abandonment was characterized by decreased pH values accompanied by decreased amounts of readily available organic carbon and nitrogen, a lower microbial biomass and decreased activity as well as a higher relative abundance of actinomycetes. The δ¹³C-signature of CO₂ indicated a preferential mineralization of grass-derived organic carbon in pasture control soils. In soils amended with grass litter the mineralization of soil organic matter was retarded (negative priming effect) and also a preferential utilization of easily available organic substances derived from the grass litter was evident. Compared to the other treatments, the pasture soil amended with grass litter showed an opposite shift in the microbial community structure towards a lower relative abundance of fungi. After addition of bracken litter to the abandoned pasture soil a positive priming effect seemed to be supported by an N limitation at the end of incubation. This was accompanied by an increase in the ratio of Gram-positive to Gram-negative bacterial PLFA marker. The differences in litter quality between grass and bracken are important triggers of changes in soil biogeochemical and soil microbial properties after land use conversion.     

The biochemical transformation of oak (Quercus robur) leaf litter consumed by the pill millipede (Glomeris marginata)

Soil macrofauna play an essential role in the initial comminution and degradation of organic matter entering the soil environment and yet the chemical effects of digestion on leaf litter are poorly understood at the molecular level. This study was undertaken to assess the selective chemical transformations that saprophagous soil invertebrates mediate in consumed leaf litter. A number of pill millipedes (Glomeris marginata) were fed oak leaves (Quercus robur) after which the biomolecular compositions (lipids and macromolecular components) of the leaves and millipede faeces were compared using a series of wet chemical techniques and subsequent analysis by gas chromatography (GC) and gas chromatography-mass spectrometry (GC/MS). It was found that the concentrations of short chain (<C₂₀) n-alkanoic acids, sterols and triacylglycerols reduced dramatically in the millipede faeces relative to the leaf litter. Hydrolysable carbohydrates and proteins both decreased in concentration in the faeces, whereas similar yields of phenolic components were observed for the cupric oxidation products of lignin, although the oxygenated functionalities were affected by passage through the millipede gut, yielding a more highly condensed state for lignin. This shows that the chemical composition of fresh organic matter entering the soil is directly controlled by invertebrates feeding upon the leaf litter and as such that they are key contributors to the early stages of diagenesis in terrestrial soils.     

Transfer of N fixed by a legume tree to the associated grass in a tropical silvopastoral system

Below-ground transfer of nitrogen (N) fixed by legume trees to associated non-N₂-fixing crops has received little attention in agroforestry, although the importance of below-ground interactions is shown in other ecosystems. We used ¹⁵N natural abundance to estimate N transfer from the legume tree Gliricidia sepium (Jacq.) Kunth ex Walp. to C₄ grass Dichanthium aristatum (Poir.) C.E. Hubb. in a silvopastoral system, where N was recycled exclusively by below-ground processes and N₂ fixation by G. sepium was the sole N input to the system. Finding a suitable reference plant, a grass without contact with tree roots or litter, was problematic because tree roots invaded adjacent grass monocrop plots and soil isotopic signature in soil below distant grass monocrops differed significantly from the agroforestry plots. Thus, we used grass cultivated under greenhouse conditions in pots filled with agroforestry soil as the reference. A model of soil ¹⁵N fractionation during N mineralization was developed for testing the reliability of that estimate. Experimental and theoretical results indicated that 9 months after greenhouse transplanting, the percentage of fixed N in the grass decreased from 35% to <1%, due to N export in cut grass and dilution of fixed N with N taken up from the soil. The effect of soil ¹⁵N fractionation on the estimate of the reference value was negligible. This indicates that potted grass is a suitable reference N transfer studies using ¹⁵N natural abundance. About one third of N in field-grown grass was of atmospheric origin in agroforestry plots and in adjacent D. aristatum grassland invaded by G. sepium roots. The concentration of fixed N was correlated with fine root density of G. sepium but not with soil isotopic signature. This suggests a direct N transfer from trees to grass, e.g. via root exudates or common mycorrhizal networks.     

Consumption of mycorrhizal and saprophytic fungi by Collembola in grassland soils

Although soil-dwelling Collembola can influence plant growth and nutrient cycling, their specific role in soil food webs is poorly understood. Soil-free microcosm studies suggest that Collembola are primarily fungivores where they feed preferentially on saprophytic fungi (SF) over other fungal types. We directly assessed collembolan consumption of arbuscular mycorrhizal fungi (AMF) and SF using plant-soil mesocosms and natural abundance stable carbon isotope techniques. Mycorrhizal Andropogon gerardii (C₄ grass) seedlings were placed in pots containing Collembola and soil from a C₃ plant dominated site, while mycorrhizal Pascopyrum smithii (C₃ grass) seedlings were placed in pots with Collembola and soil collected at a C₄ plant dominated site. After 6 weeks, collembolans assimilated carbon derived from C₃ and C₄ sources in both A. gerardii and P. smithii treatments. Comparing Collembola isotope values in AMF vs. AMF-suppressed treatments, our data show that both AMF and SF were consumed in these experimental soil environments.     

Compound-specific δC and δH analyses of plant and soil organic matter: A preliminary assessment of the effects of vegetation change on ecosystem hydrology

Here we present δ¹³C and δ²H data of long-chained, even-numbered (C₂₇-C₃₁) n-alkanes from C3 (trees) and C4 (grasses) plants and from the corresponding soils from a grassland-woodland vegetation sequence in central Queensland, Australia. Our data show that δ¹³C values of the C4 grassland species were heavier relative to those of C3 tree species from the woodland (Acacia leaves) and woody grassland (Atalaya leaves). However, n-alkanes from the C4 grasses had lighter δ²H values relative to the Acacia leaves, but showed no significant difference in δ²H values when compared with C3 Atalaya leaves. These results differ from those of previous studies, showing that C4 grasses had heavier δ²H values relative to C3 grasses and trees. Those observations have been explained by C4 plants accessing the more evaporation-influenced and isotopically heavier surface water and tree roots sourcing deeper, isotopically lighter soil water (“Two-layered soil-water system”). By comparison, our data suggest that ecosystem changes (vegetation “thickening”) can significantly alter the soil hydrological characteristics. This is shown by the heavier δ²H values in the woodland soil compared with lighter δ²H values in the grassland soil, implying that the recent vegetation change (increased tree biomass) in the woodland had altered soil hydrological conditions. Estimated δ²H values of the source-water for vegetation in the grassland and woodland showed that both trees and grasses in open settings accessed water with lighter δ²H values (avg. −46‰) compared with water accessed by trees in the woodland vegetation (avg. −7‰). These data suggest that in semi-arid environments the “two-layer” soil water concept might not apply. Furthermore, our data indicate that compound-specific δ²H and δ¹³C analyses of n-alkanes from soil organic matter can be used to successfully differentiate between water sources of different vegetation types (grasses versus trees) in natural ecosystems.     

Nature and Origin of Lipids in Clay Fractions from a Fluvisol in a Sewage Sludge Deposition Field

Few studies on free lipids in total solvent extracts from soil clay fractions directly measured by gas chromatography–mass spectrometry (GC/MS) have been reported so far. In this study, we aimed to examine the free lipids in the clay fraction separated from a Fluvisol profile on which sewage sludge was deposited 6 years ago and provide information on the sources, diagenetic processes and organic pollutants derived from the sludge. Clay fractions were separated from the four horizons of a Fluvisol and analysed for the biochemically stable lipid pool. The GC/MS analysis of the lipid fraction showed that lipid signatures were dominated by alkanes (C₁₇–C₃₃), alkanoic acids (C₁₂–C₁₈), alkanols (C₁₄–C₃₀), aromatic acids and phthalate esters. Sources of lipids show predominant bacterial contribution as shown by the alkane, fatty acids and n-alkanol distributions. The preservation of lipids of microbial origin in the clay fraction was revealed by the presence of even number, branched alkanes and short-chain and branched fatty acids. These results imply that similar pedogenic processes took place in this soil profile affected by hydromorphic conditions with some quantitative changes in the clay lipid compositions between different soil horizons. Some aromatic acids and xenobiotics such as phthalates were detected in the lipid extracts of the clay fractions in depth (0–85 cm) of the soil profile, which has implications for pollution of soils and ground waters in situations of sewage sludge deposition.     

Crop-specific differences in the concentrations of lipids in leachates from the root zone

Although lipids are involved in diverse soil processes and affect various soil properties, the contribution of rhizodeposits and the root zone to lipid concentrations and distributions in soils is unknown. For the first time, we determined the concentrations of alkanoic acids, n-alkanes and n-alkenes in root zone leachates and roots of maize and potato using gas chromatography/mass spectrometry (GC/MS). In total, the lipid concentrations of leachates were 100 μg g^?1 (maize) and 17 μg g^?1 (potato). The saturated n-alkanoic acids, ranging from n-C₁₄ to n-C₂₈ and having the maximum at n-C₂₂ (maize) and at n-C₁₆ (potato), were more abundant than the other compounds. Maize leachates had more alkanes (20 μg g^?1) than potato leachates (3.1 μg g^?1), but the members of the homologues were nearly the same. Comparison of these distributions with data for roots, microorganisms and soil indicated that the lipids in the leachates from the root zone mainly originated from abrasion of fine roots, rhizodeposits and rhizosphere microorganisms.     

Changes in soil organic carbon dynamics in an Eastern Chinese coastal wetland following invasion by a C4 plant Spartina alterniflora

The exotic C₄ grass Spartina alterniflora was intentionally introduced to tidal coastal wetlands in Jiangsu province of China in 1982. Since then it has rapidly replaced the native C₃ plant Suaeda salsa, becoming one of the dominant vegetation types in the coastal wetlands of China. Although plant invasion can change soil organic carbon (SOC) storage, little is known about how plant invasion influences C storage within soil fractions. We investigated how S. alterniflora invasion across an 8, 12 and 14-year chronosequence affected SOC and soil nitrogen (N), using soil fractionation and stable δ¹³C isotope analyses. SOC and N concentrations at 0-10 cm depth in S. alterniflora soil increased during the S. alterniflora invasion chronosequence, ranging from 3.67 to 4.90 g C kg⁻¹ soil, and from 0.307 to 0.391 g N kg⁻¹ soil. These were significantly higher than the values in the Suaeda salsa community, by 27.0-69.6% for SOC, and 21.8-55.2% for total N. The S. alterniflora-derived SOC varied from 0.40 to 0.92 g C kg⁻¹ according to mixing calculations, assuming the two possible SOC sources of S. alterniflora and S. salsa, and accounted for 10.8-18.7% of total SOC in the colonized soils. The estimated accumulative rate of SOC from C₄ (S. alterniflora) was 64.1 C kg⁻¹ soil year⁻¹ and from C₃ sources was 78.1 mg C kg⁻¹. The concentration of S. alterniflora-derived SOC significantly decreased from coarse fraction to fine fraction, and linearly increased as the period of S. alterniflora invasion increased. The highest accumulative rate of SOC from a C₄ source occurred in macroaggregates, while the highest rate from C₃ was in microaggregates. The storage of SOC derived from S. alterniflora in the macroaggregates was 0.27-0.44 g C kg⁻¹ soil, accounting for 43.1-49.1% of the total C₄derived SOC in the soil. Our results suggest that S. alterniflora invasion in coastal wetlands could facilitate SOC storage, because of the high potential for accumulation of the C which has been newly derived from S. alterniflora litter and roots.     

Thermal stability of soil organic matter pools and their δC values after C3-C4 vegetation change

Carbon isotopic composition of soils subjected to C₃-C₄ vegetation change is a suitable tool for the estimation of C turnover in soil organic matter (SOM) pools. We hypothesized that the biological availability of SOM pools is inversely proportional to their thermal stability. Soil samples from a field plot with 10.5 years of cultivation of the C₄ plant Miscanthus×gigantheus and from a reference plot under C₃ grassland vegetation were analysed by thermogravimetry coupled with differential scanning calorimetry (TG-DSC). According to differential weight losses (dTG) and energy release or consumption (DSC), five SOM pools with increasing thermal stability were distinguished: (I) 20-190 °C, (II) 190-310 °C, (III) 310-390 °C, (IV) 390-480 °C, and (V) 480-1000 °C. Their δ¹³C values were analysed by EA-IRMS. The weight losses in pool I were connected with water evaporation, since no significant C losses were measured and δ¹³C values remained unchanged. The δ¹³C of pools II and III in soil samples under Miscanthus were closer to the δ¹³C of the Miscanthus plant tissues (−11.8‰) compared to the thermally stable SOM pool V (−19.5‰). The portion of the Miscanthus-derived C₄-C in total SOM in 0-5 cm reached 55.4% in the 10.5 years. The C₄-C contribution in pool II was 60% and decreased down to 6% in pool V. The mean residence times (MRT) of SOM pools II, III, and IV were similar (11.6, 12.2, and 15.4 years, respectively), while pool V had a MRT of 163 years. Therefore, we concluded that the biological availability of thermal labile SOM pools (<480 °C) was higher, than that of the thermal stable pool decomposed above 480 °C. However, the increase of SOM stability with rising temperature was not gradual. Therefore, the applicability of the TG-DSC for the separation of SOM pools with different biological availability is limited.     

Competition for inorganic and organic N by blue oak (Quercus douglasii) seedlings, an annual grass, and soil microorganisms in a pot study

Sources of competition for limited soil resources, such as nitrogen (N), include competitive interactions among different plant species and between plants and soil microorganisms (microbes). To study these competitive interactions, blue oak seedlings (Quercus douglasii) were grown alone or grown together with an annual grass, wild oats (Avena barbata) in pots containing field soil. We injected ¹⁵N-labeled ammonium, nitrate or glycine into the soil of each pot and harvested plants 5 days later. Plant shoots and roots, soil microbial N and soil KCl-extractable N were analyzed for ¹⁵N content. When oak and grass were grown together, ¹⁵N recovery from the inorganic N treatments (NH₄⁺, or NO₃⁻) was 34, 9 and 4% for the grass, microbes and oak seedlings, respectively, and only 1% remained as KCl-extractable N. ¹⁵N recovery from the glycine treatment was 18, 22, 5% for the grass, microbes and oak seedlings, respectively, and 4% remained as KCl-extractable N. When oaks were grown alone, ¹⁵N recovery by soil microbes was 21, 48 and 40% in the NO₃⁻, NH₄⁺ and glycine treatments, respectively. N forms had no effects on ¹⁵N recovery in oak seedlings (7%) and in KCl-extractable N pool (13%). In general, total N recovery by the grass was much greater than by oaks. However, on a fine root surface area or length basis, oaks exhibited higher N uptake than the grass. Our results suggest that the high rooting density and rapid growth rate of the annual grasses such as Avena barbata made them superior competitors for available soil N when compared to blue oak seedlings and to microbes. Soil microbes were better competitors for organic than inorganic N when annual grasses were present, but preferred NH₄⁺ when competing only with oak seedlings.     

Temperature sensitivity of respiration differs among forest floor layers in a Pinus resinosa plantation

Decomposer microorganisms contribute to carbon loss from the forest floor as they metabolize organic substances and respire CO₂. In temperate and boreal forest ecosystems, the temperature of the forest floor can fluctuate significantly on a day-to-night or day-to-day basis. In order to estimate total respiratory CO₂ loss over even relatively short durations, therefore, we need to know the temperature sensitivity (Q₁₀) of microbial respiration. Temperature sensitivity has been calculated for microbes in different soil horizons, soil fractions, and at different depths, but we would suggest that for some forests, other ecologically relative soil portions should be considered to accurately predict the contribution of soil to respiration under warming. The floor of many forests is heterogeneous, consisting of an organic horizon comprising a few more-or-less distinct layers varying in decomposition status. We therefore determined at various measurement temperatures the respiration rates of litter, F-layer, and H-layer collected from a Pinus resinosa plantation, and calculated Q₁₀ values for each layer. Q₁₀ depended on measurement temperature, and was significantly greater in H-layer than in litter or F-layer between 5 and 17 °C. Our results indicate, therefore, that as the temperature of the forest floor rises, the increase in respiration by the H-layer will be disproportionate to the increase by other layers. However, change in respiration by the H-layer associated with change in temperature may contribute minimally or significantly to changes of total forest floor respiration in response to changes in temperature depending on the depth and thickness of the layer in different forest ecosystems.     

Exotic plant influences soil nematode communities through litter input

The impact of exotic plant invasions on soil communities and nutrient cycling processes has received an increasing attention in recent years. To test whether the exotic plant invasions affect nematode communities through altering litter quality, we compared mass loss and nematode colonization during the stem litter decomposition of invasive Spartina alterniflora and native Phragmites australis in salt marshes of the Yangtze River estuary, China. Plastic drinking straws were synchronously used as controls. The addition of plant residues was found stimulating the growth of nematodes, particularly bacterial feeders on day 16 after burial. A top-down control of bacterivous nematodes by carnivores existed in nematode succession during the litter decomposition. With higher nitrogen content and lower C:N ratio, stem litter of the invasive S. alterniflora decayed faster and supported more abundant nematodes than the native P. australis. The greater nematode abundance in S. alterniflora was mainly due to two dominant genera of bacterial nematodes, namely Diplolaimelloides and Diplolaimella. Lower values of maturity index and structure index in S. alterniflora than in P. australis litter indicate that a more degraded food web condition resulted from the faster litter decay. A considerable difference in nematode community structures between two litter types only occurred in a certain period of the decomposition (from 8 to 32 days after burial), suggesting that the changes in faunal community structure are time dependent. In summary, this study confirmed the hypothesis that the invasion of S. alterniflora stimulates the growth of bacterial nematodes by producing higher quality of litter than the native P. australis. The results obtained here suggest that the invasion of exotic plant is likely to alter ecosystem functions indirectly through exerting its effect on soil decomposer communities such as nematodes.     

不同生境条件下西藏原始冷杉林凋落物分解特征与土壤养分的关系

[目的]研究不同生境条件下(林内、林外、林缘)藏东南急尖长苞冷杉林(Abies georgei var.smithii)凋落物分解特征与土壤养分特征之间的关系,为深入了解高寒高山森林生态系统物质循环过程提供依据。[方法]采用野外分解袋法和室内分析相结合,在林内、林外、林缘3种不同生境条件下对藏东南急尖长苞冷杉林凋落物进行了原位分解试验。[结果]分解速率总体上呈现出:林内林缘林外的特点,逐月分解率的变异系数表现为:林内(34.83%)林缘(57.35%)林外(72.09%);Olson指数衰减模型的模拟结果显示不同生境条件下(林内、林缘、林外)凋落物分解50%需要的时间为2.11,2.52,2.34 a,分解95%需要的时间为8.96,10.01,10.84 a;3种不同生境土壤养分在空间上差异显著,林内生境中与凋落物分解速率呈现极显著相关的土壤养分因子有土壤总有机碳(TOC)含量、N含量、土壤微生物量碳(SMBC)含量、土壤微生物量氮(SMBN)含量以及W_C∶W_N值;林外、林缘生境中与凋落物分解速率相关性最大的为土壤TOC含量,其次为W_C∶W_N值。[结论]生境条件的差异对凋落物分解速率有显著影响,在不同的生境条件下对凋落物分解影响起主导作用的土壤养分因素不同,凋落物—土壤生物地化循环紧密联系,相互作用关系复杂,生境作用效应突出。     

Short-term C4 plant Spartina alterniflora invasions change the soil carbon in C3 plant-dominated tidal wetlands on a growing estuarine Island

Spartina alterniflora is an invasive C₄ perennial grass, native to North America, and has spread rapidly along the east coast of China since its introduction in 1979. Since its intentional introduction to the Jiuduansha Island in the Yangtze River estuary, Spartina alterniflora community has become one of the dominant vegetation types. We investigated the soil carbon in the Spartina alterniflora community and compared it with that of the native C₃Scirpus mariqueter community by measuring total soil carbon (TC), soil organic carbon (SOC), total soil nitrogen (TN), and the stable carbon isotope composition (δ¹³C) of various fractions. TC and SOC were significantly higher in Spartina alterniflora in the top 60 cm of soil. However, there was no significant difference in soil inorganic carbon (IC) between the two communities. Stable carbon isotopic analysis suggests that the fraction of SOC pool contributed by Spartina alterniflora varied from 0.90% to 10.64% at a soil depth of 0-100 cm with a greater percentage between 20 and 40 cm deep soils. The δ¹³C decreased with increasing soil depth in both communities, but the difference in δ¹³C among layers of the top 60 cm soil was significant (p<0.05), while that for the deeper soil layers (>60 cm) was not detected statistically. The changes in δ¹³C with depth appeared to be associated with the small contribution of residues from Spartina alterniflora at greater soil depth that was directly related to the vertical root distribution of the species.     

Rhizosphere activity, grass species and N availability effects on the soil C and N cycles

We examined whether grass species and soil nitrogen (N) availability could enhance Carbon (C) and N turnover during root litter decay in grassland. Three species with increasing competitiveness (Festuca ovina, Dactylis glomerata and Lolium perenne) were grown at two N fertiliser levels in an undisturbed grassland soil, in which soil organic fractions derived for the last 9 years from Lolium root litter which was ¹³C-depleted. During the subsequent experimental year, the C turnover was calculated using the respective δ¹³C values of the old and new C in the root phytomass, in two Particulate Organic Matter (POM) fractions above 200 μm and in the lightest part of the aggregated soil fraction between 50 and 200 μm. Soil N availability was monitored during the regrowth periods with ion exchange resins (IER). The C decay rates of each particle size fraction were calculated with a simple mechanistic model of C dynamics. The N mineralisation immobilisation turnover (MIT) was characterised by dilution of ¹⁵N-labelled fertiliser in the N harvestThe C:N ratio and the residence time of C in the fractions decreased with particle size. The presence of a grass rhizosphere increased the decay rate of old C. Accumulation of new C in particle size fractions increased with species competitiveness and with N supply. Species competitiveness increased C turnover in the aggregated fraction, as a result of greater accumulation of new C and faster decay of old C. Fertiliser N increased N turnover and C mineralisation in the SOM. Species competitiveness decreased soil -N exchanged with the IER and increased dissolved organic C (DOC) content. The nature of the current rhizosphere is thus an important factor driving C and N transformations of the old root litter, in relation with grass species strategy. Plant competitiveness may stimulate the C and N turnover in the more evolved SOM fractions in a similar way to the mineral N supply.     

Grasses, litter, and their interaction affect microbial biomass and soil enzyme activity

Plant effects on ecosystem processes are mediated through plant-microbial interactions belowground and soil enzyme assays are commonly used to directly relate microbial activity to ecosystem processes. Live plants influence microbial biomass and activity via differences in rhizosphere processes and detrital inputs. I utilized six grass species of varying litter chemistry in a factorial greenhouse experiment to evaluate the relative effect of live plants and detrital inputs on substrate-induced respiration (SIR, a measure of active microbial biomass), basal respiration, dissolved organic carbon (DOC), and the activities of β-glucosidase, β-glucosaminidase, and acid phosphatase. To minimize confounding variables, I used organic-free potting media, held soil moisture constant, and fertilized weekly. SIR and enzyme activities were 2-15 times greater in litter-addition than plant-addition treatments. Combining live plants with litter did not stimulate microbial biomass or activity above that in litter-only treatments, and β-glucosidase activity was significantly lower. Species-specific differences in litter N (%) and plant biomass were related to differences in β-glucosaminidase and acid phosphatase activity, respectively, but had no apparent effect on β-glucosidase, SIR, or basal respiration. DOC was negatively related to litter C:N, and positively related to plant biomass. Species identity and living plants were not as important as litter additions in stimulating microbial activity, suggesting that plant effects on soil enzymatic activity were driven primarily by detrital inputs, although the strength of litter effects may be moderated by the effect of growing plants.     

Competition between invasive earthworms (Amynthas corticis, Megascolecidae) and native North American millipedes (Pseudopolydesmus erasus, Polydesmidae): Effects on carbon cycling and soil structure

Invasive earthworms can have significant impacts on C dynamics through their feeding, burrowing, and casting activities, including the protection of C in microaggregates and alteration of soil respiration. European earthworm invasion is known to affect soil micro- and mesofauna, but little is known about impacts of invasive earthworms on other soil macrofauna. Asian earthworms (Amynthas spp.) are increasingly being reported in the southern Appalachian Mountains in southeastern North America. This region is home to a diverse assemblage of native millipedes, many of which share niches with earthworm species. This situation indicates potential for earthworm-millipede competition in areas subject to Amynthas invasion.In a laboratory microcosm experiment, we used two ¹³C enriched food sources (red oak, Quercus rubra, and eastern hemlock, Tsuga canadensis) to assess food preferences of millipedes (Pseudopolydesmus erasus), to determine the effects of millipedes and earthworms (Amynthas corticis) on soil structure, and to ascertain the nature and extent of the interactions between earthworms and millipedes. Millipedes consumed both litter species and preferred red oak litter over eastern hemlock litter. Mortality and growth of millipedes were not affected by earthworm presence during the course of the experiment, but millipedes assimilated much less litter-derived C when earthworms were present.Fauna and litter treatments had significant effects on soil respiration. Millipedes alone reduced CO₂ efflux from microcosms relative to no fauna controls, whereas earthworms alone and together with millipedes increased respiration, relative to the no fauna treatment. CO₂ derived from fresh litter was repressed by the presence of macrofauna. The presence of red oak litter increased CO₂ efflux considerably, compared to hemlock litter treatments.Millipedes, earthworms, and both together reduced particulate organic matter. Additionally, earthworms created significant shifts in soil aggregates from the 2000-250 and 250-53 μm fractions to the >2000 μm size class. Earthworm-induced soil aggregation was lessened in the 0-2 cm layer in the presence of millipedes. Earthworms translocated litter-derived C to soil throughout the microcosm.Our results suggest that invasion of ecosystems by A. corticis in the southern Appalachian Mountains is unlikely to be limited by litter species and these earthworms are likely to compete directly for food resources with native millipedes. Widespread invasion could cause a net loss of C due to increased respiration rates, but this may be offset by C protected in water-stable soil aggregates.     

Carbon dioxide emissions of soils under pure and mixed stands of beech and spruce, affected by decomposing foliage litter mixtures

Soil respiration is the largest terrestrial source of CO₂ to the atmosphere. In forests, roughly half of the soil respiration is autotrophic (mainly root respiration) while the remainder is heterotrophic, originating from decomposition of soil organic matter. Decomposition is an important process for cycling of nutrients in forest ecosystems. Hence, tree species induced changes may have a great impact on atmospheric CO₂ concentrations. Since studies on the combined effects of beech-spruce mixtures are very rare, we firstly measured CO₂ emission rates in three adjacent stands of pure spruce (Picea abies), mixed spruce-beech and pure beech (Fagus sylvatica) on three base-rich sites (Flysch) and three base-poor sites (Molasse; yielding a total of 18 stands) during two summer periods using the closed chamber method. CO₂ emissions were higher on the well-aerated sandy soils on Molasse than on the clayey soils on Flysch, characterized by frequent water logging. Mean CO₂ effluxes increased from spruce (41) over the mixed (55) to the beech (59) stands on Molasse, while tree species effects were lower on Flysch (30-35, mixed > beech = spruce; all data in mg CO₂-C m⁻² h⁻¹). Secondly, we studied decomposition after fourfold litter manipulations at the 6 mixed species stands: the O_i - and O_e horizons were removed and replaced by additions of beech -, spruce - and mixed litter of the adjacent pure stands of known chemical quality and one zero addition (blank) in open rings (20 cm inner diameter), which were covered with meshes to exclude fresh litter fall. Mass loss within two years amounted to 61-68% on Flysch and 36-44% on Molasse, indicating non-additive mixed species effects (mixed litter showed highest mass loss). However, base cation release showed a linear response, increasing from the spruce - over the mixed - to the beech litter. The differences in N release (immobilization) resulted in a characteristic converging trend in C/N ratios for all litter compositions on both bedrocks during decomposition. In the summers 2006 and 2007 we measured CO₂ efflux from these manipulated areas (a closed chamber fits exactly over such a ring) as field indicator of the microbial activity. Net fluxes (subtracting the so-called blank values) are considered an indicator of litter induced changes only and increased on both bedrocks from the spruce - over the mixed - to the beech litter. According to these measurements, decomposing litter contributed between 22-32% (Flysch) and 11-28% (Molasse) to total soil respiration, strengthening its role within the global carbon cycle.     

Contribution of primary organic matter to the fatty acid pool in agricultural soils

Fatty acids as major compounds of soil lipids may affect many soil properties, but the input and turnover rates in soil are largely unknown. The objective of this study was to identify and quantify fatty acids in soils as a result of input from primary sources such as plant residues, farmyard manure and soil organisms, and to evaluate the corresponding turnover- and stabilization processes. The concentrations of n-C_10:0 to n-C_34:0 fatty acids were determined in the Ap horizon of a Phaeozem with long-term cropping of rye and maize and the treatments ‘Unfertilized’ (‘U’) and fertilized with ‘Farmyard manure’ (‘FYM’). The most important primary sources of fatty acids such as rye and maize stubble and roots, soil micro- and mesofauna, and the applied FYM were also investigated. The quantification of fatty acids by gas chromatography/mass spectrometry (GC/MS) showed that long-term FYM application led to larger concentrations of n-alkyl fatty acids in the plots grown with rye (‘U’: 48.1 μg g⁻¹, ‘FYM’: 57.7 μg g⁻¹, **P≤0.01, n=3) and maize (‘U’: 17.0 μg g⁻¹, ‘FYM’: 23.4 μg g⁻¹, ***P≤0.001, n=3). The observed bimodal fatty acid distribution in soils from n-C_10:0 to n-C_21:0 and from n-C_21:0 to n-C_34:0 with a predominance at n-C_16:0 and at n-C_28:0 was apparently due to input from crop residues, soil organisms and FYM. The short-chain lengths may have originated from the investigated primary sources. The major contributors to the long-chain lengths, with a maximum at n-C_28:0, were rye stubble and FYM. A change in mono-culture from rye to maize, 38 years prior to sampling, led to a decrease in fatty acid concentrations by factors of about 2.8 (‘U’) and 2.5 (‘FYM’). Therefore, rye-derived fatty acids and soil tillage had a larger impact on fatty acid pools than the input of primary organic matter. The changes in fatty acid distributions and pools under the consideration of the quantified input of primary organic matter led to the conclusion that the short-chained fatty acids were more rapidly decomposed than the long-chains.     

Impacts of extreme winter warming events on litter decomposition in a sub-Arctic heathland

Arctic climate change is expected to lead to a greater frequency of extreme winter warming events. During these events, temperatures rapidly increase to well above 0 °C for a number of days, which can lead to snow melt at the landscape scale, loss of insulating snow cover and warming of soils. However, upon return of cold ambient temperatures, soils can freeze deeper and may experience more freeze-thaw cycles due to the absence of a buffering snow layer. Such loss of snow cover and changes in soil temperatures may be critical for litter decomposition since a stable soil microclimate during winter (facilitated by snow cover) allows activity of soil organisms. Indeed, a substantial part of fresh litter decomposition may occur in winter. However, the impacts of extreme winter warming events on soil processes such as decomposition have never before been investigated. With this study we quantify the impacts of winter warming events on fresh litter decomposition using field simulations and lab studies.Winter warming events were simulated in sub-Arctic heathland using infrared heating lamps and soil warming cables during March (typically the period of maximum snow depth) in three consecutive years of 2007, 2008, and 2009. During the winters of 2008 and 2009, simulations were also run in January (typically a period of shallow snow cover) on separate plots. The lab study included soil cores with and without fresh litter subjected to winter-warming simulations in climate chambers.Litter decomposition of common plant species was unaffected by winter warming events simulated either in the lab (litter of Betula pubescens ssp. czerepanovii), or field (litter of Vaccinium vitis-idaea, and B. pubescens ssp. czerepanovii) with the exception of Vaccinium myrtillus (a common deciduous dwarf shrub) that showed less mass loss in response to winter warming events. Soil CO₂ efflux measured in the lab study was (as expected) highly responsive to winter warming events but surprisingly fresh litter decomposition was not. Most fresh litter mass loss in the lab occurred during the first 3-4 weeks (simulating the period after litter fall).In contrast to past understanding, this suggests that winter decomposition of fresh litter is almost non-existent and observations of substantial mass loss across the cold season seen here and in other studies may result from leaching in autumn, prior to the onset of “true” winter. Further, our findings surprisingly suggest that extreme winter warming events do not affect fresh litter decomposition.