Nitrogen losses from two grassland soils with different fungal biomass

Nitrogen losses from agricultural grasslands cause eutrophication of ground- and surface water and contribute to global warming and atmospheric pollution. It is widely assumed that soils with a higher fungal biomass have lower N losses, but this relationship has never been experimentally confirmed. With the increased interest in soil-based ecosystem services and sustainable management of soils, such a relationship would be relevant for agricultural management. Here we present a first attempt to test this relationship experimentally. We used intact soil columns from two plots from a field experiment that had consistent differences in fungal biomass (68 ± 8 vs. 111 ± 9 μg C g⁻¹) as a result of different fertilizer history (80 vs. 40 kg N ha⁻¹ y⁻¹ as farm yard manure), while other soil properties were very similar. We performed two greenhouse experiments: in the main experiment the columns received either mineral fertilizer N or no N (control). We measured N leaching, N₂O emission and denitrification from the columns during 4 weeks, after which we analyzed fungal and bacterial biomass and soil N pools. In the additional ¹⁵N experiment we traced added N in leachates, soil, plants and microbial biomass. We found that in the main experiment, N₂O emission and denitrification were lower in the high fungal biomass soil, irrespective of the addition of fertilizer N. Higher ¹⁵N recovery in the high fungal biomass soil also indicated lower N losses through dentrification. In the main experiment, N leaching after fertilizer addition showed a 3-fold increase compared to the control in low fungal biomass soil (11.9 ± 1.0 and 3.9 ± 1.0 kg N ha⁻¹, respectively), but did not increase in high fungal biomass soil (6.4 ± 0.9 after N addition vs. 4.5 ± 0.8 kg N ha⁻¹ in the control). Thus, in the high fungal biomass soil more N was immobilized. However, the ¹⁵N experiment did not confirm these results; N leaching was higher in high fungal biomass soil, even though this soil showed higher immobilization of ¹⁵N into microbial biomass. However, only 3% of total ¹⁵N was found in the microbial biomass 2 weeks after the mineral fertilization. Most of the recovered ¹⁵N was found in plants (approximately 25%) and soil organic matter (approximately 15%), and these amounts did not differ between the high and the low fungal biomass soil. Our main experiment confirmed the assumption of lower N losses in a soil with higher fungal biomass. The additional ¹⁵N experiment showed that higher fungal biomass is probably not the direct cause of higher N retention, but rather the result of low nitrogen availability. Both experiments confirmed that higher fungal biomass can be considered as an indicator of higher nitrogen retention in soils.     

Soil animals and ecosystem processes: How much does nutrient cycling explain?

Trophic-dynamic hypotheses have been extensively tested by manipulating the presence of soil animals in experimental laboratory microcosms. Soil animals typically have pronounced effects on microbial populations, nutrient cycling and plant growth. However, because often only the total effect has been reported, the relative importance of feeding interactions versus non-trophic effects remains obscure. Using simple calculations based on mass conservation I argue that the observed faunal effect on microbes and system functioning is often larger than can be explained by trophic dynamics and nutrient cycling. Non-trophic effects may help to explain why microcosm experiments have failed to support trophic-dynamic predictions like trophic cascades. Since such effects are also likely independent of species identity and population density, they may facilitate the interpretation of experiments where decomposition processes have been found to be largely insensitive to soil fauna diversity.     

秸秆还田下氮肥用量对稻田养分淋洗的影响

通过田间试验,研究秸秆还田配施氮肥对稻田土壤养分淋洗的影响。结果表明,随氮肥用量增加,田间渗漏水中NH4+-N、NO3--N、全氮浓度随之增加;与秸秆未还田相比,秸秆还田降低了田面水与渗漏水中NH4+-N、NO3--N的浓度;秸秆还田下各处理30cm土层渗漏水中全氮和NO3--N浓度最高,其浓度范围分别为1.09~12.76mg·L-1和0.76~3.74mg·L-1;全磷浓度范围为0.02~0.79mg·L-1,田面水中全磷浓度随施氮量增加而增加,30cm渗漏水中全磷浓度大于60cm渗漏水。氮肥用量180kg·hm-2时,施肥后5~10d内30cm、60cm渗漏水中的养分以NH4+-N为主,其后均以NO3--N为主。氮肥与秸秆配合施用,可降低田面水和渗漏水中的氮磷浓度,改善肥料利用效率。     

Biological and physical effects of non-native earthworms on nitrogen cycling in riparian soils

Global nitrogen cycling is being altered by anthropogenic disturbances including invasion by non-native species. European and Asian earthworms have invaded northern temperate forests in North America with dramatic consequences for litter thickness, forest floor plant diversity, and soil nitrogen cycling. Invasive earthworms present at the boundary of terrestrial and aquatic ecosystems (i.e., riparian zones) may alter the flux of nitrogen into adjacent aquatic ecosystems. We examined how nitrogen cycling in riparian soil responds to amendments of invasive earthworms or artificial earthworm burrows. In earthworm-free riparian plots (0.25 m²), we established treatments of invasive earthworms (60 g fresh mass·m⁻²), artificial burrows (120 m⁻²), or control plots and sampled the plots after 30 days. Before and after treatment application we measured major soil characteristics (water-filled pore space, organic matter, and pH), nitrogen pools (exchangeable NH₄⁺ and NO₃⁻), and nitrogen transformation rates (net N-mineralization, net nitrification, and denitrification). Exchangeable NH₄⁺ and NO₃⁻ changed through time but did not differ among treatments. Net N-mineralization and net nitrification rates did not change through time and were similar across all treatments. However, denitrification rates in plots with added earthworms were 4 times greater than rates in control and burrow-only plots, which represents a large rapid increase in gaseous nitrogen flux out of these riparian soils. For all response variables, artificial burrows responded similarly to control plots, suggesting that earthworm biological activity (i.e., feeding, excretion, and mucus production) rather than physical effects (i.e., burrowing and soil aeration) drove the changes in nitrogen cycling. Examination of soil nitrogen pool and flux measurements suggest that this increase in denitrification was coupled with NH₄⁺ consumption by nitrifying bacteria, but future studies are needed to confirm this hypothesis. We conclude that the activity of invasive earthworms in riparian zones can increase the flux of N out of riparian zones, but the hydrologic context of the riparian zone (e.g., pore-water residence time) ultimately controls whether denitrification or nitrate leaching is the dominant flux of N.     

Modeling nitrogen mineralization at surface and deep layers of sandy soils

We evaluated potential soil nitrogen mineralization of 46 sandy fields of the Pampas for determining the contribution of deep layers to mineralization and modeling its trend in depth as a possible tool for improving current existing mineralization models based on surface data. Mineralization, total and mineral nitrogen decreased with depth. A potential model fitted well to these variables (R² = 0.95–0.99), but mineralization showed a more stratified profile. Consequently, the fraction of total nitrogen mineralized decreased with depth despite soils had constant texture across the profile. Potential mineralization to 1 m depth could be estimated using data from the 0–0.2-m soil layer and the average curvature of the potential model (R² = 0.60) or linear regression methods (R² = 0.71). Another estimation of potential mineralization could be performed by developing a pedotransfer function which used as predictors total nitrogen and depth (R² = 0.62), without the need of laboratory incubations. Our results showed that for sandy soils, deep nitrogen mineralization account for 40% of soil mineralization and can be assessed using surface data or the total nitrogen content of the soils. Because surface soil mineralization and whole profile mineralization were highly correlated, it is improbable that field mineralization modeling may be improved using deep data in these soils.     

Protein breakdown represents a major bottleneck in nitrogen cycling in grassland soils

Proteins represent the dominant input of organic N into most ecosystems and they also constitute the largest store of N in soil organic matter. The extracellular protease mediated breakdown of proteins to amino acids therefore represents a key step regulating N cycling in soil. In this study we investigated the influence of a range of environmental factors on the rate of protein mineralization in a grazed grassland and fallow agricultural soil. The protein turnover rates were directly compared to the rates of amino acid mineralization under the same conditions. Uniformly ¹⁴C-labelled soluble protein and amino acids were added to soil and the rate of ¹⁴CO₂ evolution determined over 30 d. Our results indicate that the primary phase of protein mineralization was approximately 20 ± 3 fold slower that the rate of amino acid mineralization. The addition of large amounts of inorganic NO₃⁻ and NH₄⁺ to the soil did not repress the rate of protein mineralization suggesting that available N does not directly affect protease activity in the short term. Whilst protein mineralization was strongly temperature sensitive, the presence of plants and the addition of humic and tannic acids had relatively little influence on the rate of soluble protein degradation in this fertile grassland soil. Our results suggests that the extracellular protease mediated cleavage of proteins to amino acids rather than breakdown of amino acids to NH₄⁺ represents the limiting step in soil N cycling.     

Patterns in the aggregate stability of Mancos Shale derived soils

Salt and sediment contributions to the Colorado River and its tributaries pose economic and environmental concerns for the United States and Mexico. Land use decisions promoting the aggregation of Mancos Shale derived soils are one way to reduce the transportation of salts and sediments to water resources. We used a simple field test of soil aggregate stability to determine the site characteristics influencing the soil stability of sedimentary marine shale in the Gunnison Gorge National Conservation Area in Southwestern Colorado. Ninety-six 1 m² plots were intensively sampled to explore relationships between soil stability and the biological, chemical and physical site characteristics.     

The response of organic matter mineralisation to nutrient and substrate additions in sub-arctic soils

Global warming in the Arctic may alter decomposition rates in Arctic soils and therefore nutrient availability. In addition, changes in the length of the growing season may increase plant productivity and the rate of labile C input below ground. We carried out an experiment in which inorganic nutrients (NH₄NO₃ and NaPO₄) and organic substrates (glucose and glycine) were added to soils sampled from across the mountain birch forest-tundra heath ecotone in northern Sweden (organic and mineral soils from the forest, and organic soil only from the heath). Carbon dioxide production was then monitored continuously over the following 19 days. Neither inorganic N nor P additions substantially affected soil respiration rates when added separately. However, combined N and P additions stimulated microbial activity, with the response being greatest in the birch forest mineral soil (57% increase in CO₂ production compared with 26% in the heath soil and 8% in the birch forest organic soil). Therefore, mineralisation rates in these soils may be stimulated if the overall nutrient availability to microbes increases in response to global change, but N deposition alone is unlikely to enhance decomposition. Adding either, or both, glucose and glycine increased microbial respiration. Isotopic separation indicated that the mineralisation of native soil organic matter (SOM) was stimulated by glucose addition in the heath soil and the forest mineral soil, but not in the forest organic soil. These positive ‘priming’ effects were lost following N addition in forest mineral soil, and following both N and P additions in the heath soil. In order to meet enhanced microbial nutrient demand, increased inputs of labile C from plants could stimulate the mineralisation of SOM, with the soil C stocks in the tundra-heath potentially most vulnerable.     

Tillage and cropping sequence impacts on nitrogen cycling in dryland farming in eastern Montana, USA

Information on N cycling in dryland crops and soils as influenced by long-term tillage and cropping sequence is needed to quantify soil N sequestration, mineralization, and N balance to reduce N fertilization rate and N losses through soil processes. The 21-yr effects of the combinations of tillage and cropping sequences was evaluated on dryland crop grain and biomass (stems + leaves) N, soil surface residue N, soil N fractions, and N balance at the 0–20 cm depth in Dooley sandy loam (fine-loamy, mixed, frigid, Typic Argiboroll) in eastern Montana, USA. Treatments were no-tilled continuous spring wheat (Triticum aestivum L.) (NTCW), spring-tilled continuous spring wheat (STCW), fall- and spring-tilled continuous spring wheat (FSTCW), fall- and spring-tilled spring wheat–barley (Hordeum vulgare L.) (1984–1999) followed by spring wheat–pea (Pisum sativum L.) (2000–2004) (FSTW-B/P), and spring-tilled spring wheat–fallow (STW-F). Nitrogen fractions were soil total N (STN), particulate organic N (PON), microbial biomass N (MBN), potential N mineralization (PNM), NH₄-N, and NO₃-N. Annualized crop grain and biomass N varied with treatments and years and mean grain and biomass N from 1984 to 2004 were 14.3–21.2 kg N ha⁻¹ greater in NTCW, STCW, FSTCW, and FSTW-B/P than in STW-F. Soil surface residue N was 9.1–15.2 kg N ha⁻¹ greater in other treatments than in STW-F in 2004. The STN at 0–20 cm was 0.39–0.96 Mg N ha⁻¹, PON 0.10–0.30 Mg N ha⁻¹, and PNM 4.6–9.4 kg N ha⁻¹ greater in other treatments than in STW-F. At 0–5 cm, STN, PON, and MBN were greater in STCW than in FSTW-B/P and STW-F. At 5–20 cm, STN and PON were greater in NTCW and STCW than in STW-F, PNM and MBN were greater in STCW than in NTCW and STW-F, and NO₃-N was greater in FSTW-B/P than in NTCW and FSTCW. Estimated N loss through leaching, volatilization, or denitrification at 0–20 cm depth increased with increasing tillage frequency or greater with fallow than with continuous cropping and ranged from 9 kg N ha⁻¹ yr⁻¹ in NTCW to 46 kg N ha⁻¹ yr⁻¹ in STW-F. Long-term no-till or spring till with continuous cropping increased dryland crop grain and biomass N, soil surface residue N, N storage, and potential N mineralization, and reduced N loss compared with the conventional system, such as STW-F, at the surface 20 cm layer. Greater tillage frequency, followed by pea inclusion in the last 5 out of 21 yr in FSTW-B/P, however, increased N availability at the subsurface layer in 2004.     

甘肃省农田氮磷钾养分平衡状况探析

对甘肃省农田N、P、K养分平衡状况分析结果表明,甘肃省农田N、P、K养分平衡状况为N、P盈余,而K素亏损。     

阿特拉津在土壤, 矿物质及堆肥中的吸附, 运输和转化

Atrazine is a widely used herbicide for controlling weeds on both agricultural and nonagricultural land,which is equally detected in water supplies beyond safe concentrations.Although the presence of atrazine metabolites is an indication of herbicide degradation,some of them still exhibit toxicity,greater water solubility and weaker interaction with soil components than atrazine.Hence,studies with atrazine in the environment are of interest because of its potential to contaminate drinking water sources.Data on atrazine availability for transport,plant uptake,and microbial degradation and mineralization are therefore required to perform more comprehensive and realistic environmental risk assessments of its environmental fate.This review presents an account of the sorption-desorption phenomenon of atrazine on soil and other sorbents by revisiting the several mechanisms of atrazine-sorbent binding reported in the literature.The retention and transport of atrazine in soils;the influence of organic matter on atrazine sorption;the interactions of atrazine with humic substances,atrazine uptake by plants,atrazine bioccumulation and microbial degradation;atrazine transformation in composting environments;and finally atrazine removal by biosorption are discussed.     

草原土壤：分布、分类与演化

草原土壤系指草原、湿草原和森林草原植被下形成的富含有机质、盐基饱和度高的暗色土壤.世界上此类土壤面积约900万km2,我国约76万km2,是粮食生产的重要基地.本文比较了国际上草原土壤分类.不论俄罗斯或中国,均腐土开垦以后土壤有机质开始下降迅速,而后减缓,逐步达到与当地生物气候相适应的水平.所以至今仍保持一定厚度的黑土层.在均腐土资源的利用和保护上,一方面应保持其有机质平衡,另一方面更重要的是严防水土流失,以保证其生产能力.     

Implications of Azospirillum brasilense inoculation and nutrient addition on maize in soils of the Brazilian Cerrado under greenhouse and field conditions

The combination of nitrogen and Azospirillum can ensure greater nutrient absorption and crop yield in agricultural areas using high technology. Thus, the objective was to evaluate maize response to Azopirillum brasilense (AZ) inoculation and nutrient (macronutrients and micronutrients) application under greenhouse and field conditions in clay and sandy soils of the Brazilian Cerrado. In the greenhouse assays, the following parameters were measured: shoot dry weight (SDW), root dry weight (RDW), and root volume (RV). In the field experiments, the maize yield was determined after drying the grains at 60 °C for 48 h. In clay soil, there was a significant increase in the SDW, RDW and RV in the treatment with AZ concentrated (10¹¹ cells ml⁻¹ of inoculum) when compared with the control treatment and the treatment with AZ diluted (10⁶ cells ml⁻¹ of inoculum). In this soil, adding micronutrients did not affect the maize response under greenhouse conditions. In sandy soil, there was no difference between the AZ treatment and the control, except for treatments where nutrients and AZ were both added leading to a significant increase in the maize response. In both soils, the RV:RDW ratio was higher in the treatment with AZ concentrated compared to that in the treatment with AZ diluted, but the yield response depended on the addition of nutrients. Inoculation with A. brasilense gave comparable yield to the nitrogen treatment. The grain production was increased by 29% in the treatment with A. brasilense and nitrogen compared to nitrogen fertilization alone. In this study, the yield response was affected significantly when maize was inoculated with A. brasilense, but this response was dependent on the soil type under greenhouse conditions.     

Soil respiration and methane flux in adjacent forest,grassland, and cornfield soils in Hokkaido,Japan

Soil respiration and methane flux from adjacent forest, grassland, and cornfield were measured by using the closed chamber method from June to November, 1999 in Shizunai, Hokkaido, Japan, where the soil was an Aquic Humic Udivitrands derived from volcanic ash. The forest soil absorbed methane, at arate ranging from -0.12 to -0.02 mg C m_-2 h_-1, while the grassland soil emitted methane, at a rate ranging from undetectable levels to 0.18 mg C m_-2 h_-1. In the cornfield soil methane flux ranged from -0.01 to 0.04 mg C m_-2 h_-1. The soil respiration rate varied from 3 to 230 mg C m_-2 h_-1, 27 to 372 mg C m_-2 h_-1, and 29 to 156 mg C m_-2 h_-1 for the cornfield, grassland, and forest soils, respectively. Linear regression analysis demonstrated that the methane flux rate was positively correlated with the soil water-filled pore space (WFPS), and negatively correlated with the relative gas diffusion coefficient (D/D _o) and air-filled pore space (AFPS). Soil respiration rates were positively correlated with the soil temperature at all the sites. The Q ₁₀ value was 4.8, 3.3, and 1.9 for the cornfield, grassland, and forest soils, respectively.     

华北平原典型农田CO2和N2O排放通量及其与土壤养分动态和施肥的关系

对华北平原小麦-棉花(麦棉)、小麦-大豆(麦豆)、小麦-玉米(麦玉)轮作田的CO2和N2O排放通量进行了测定,分析了温室气体排放通量与土壤中碳、氮元素、气温以及施肥等之间的关系。主要结论:1)麦棉、麦豆、麦玉田的土壤CO2平均排放通量分别为CO2-C 141.7、109.8、128.2 mg.m-2.h-1,其中夏播作物的排放通量高于小麦季;2)麦棉、麦豆及麦玉田作物生长季的土壤N2O平均排放通量分别为N2O-N 98.8、38.9、44.7μg.m-2.h-1,也表现为麦后季作物的排放量高于小麦季;3)同一生育期中不同处理的N2O排放主要与土壤中无机氮含量相关,不同生育期的N2O排放通量主要受不同生育期的土壤温度及水分状况的影响;4)在施肥灌溉后的9 d内土壤N2O排放通量较高,之后逐渐降低,至施肥后22～27 d即与不施肥处理的排放持平。     

Degradation processes and nutrient constraints in sodic soils

Accumulation of excess sodium (Na⁺) in a soil causes numerous adverse phenomena, such as changes in exchangeable and soil solution ions and soil pH, destabilization of soil structure, deterioration of soil hydraulic properties, and increased susceptibility to crusting, runoff, erosion and aeration, and osmotic and specific ion effects on plants. In addition, serious imbalances in plant nutrition usually occur in sodic soils, which may range from deficiencies of several nutrients to high levels of Na⁺. The structural changes and nutrient constraints in such soils ultimately affect crop growth and yield. The principal factor that determines the extent of adverse effects of Na⁺ on soil properties is the accompanying electrolyte concentration in the soil solution, with low concentration promoting the deleterious effects of exchangeable Na⁺ even at exchangeable sodium percentage (ESP) levels less than 5. Consequent to an increase in the use of poor quality waters and soils for crop production, the problems of sodic soils can be expected to increase in future. The mechanisms that explain sodic behaviour can provide a framework in which slaking, swelling and dispersion of clay together with nutrient constraints in sodic soils may be assessed so that the practices to manage such soils can be refined for long‐term sustainable agriculture. Copyright © 2002 John Wiley & Sons, Ltd.     

Cattle grazing drives nitrogen and carbon cycling in a temperate salt marsh

We examined the impact of long-term cattle grazing on soil processes and microbial activity in a temperate salt marsh. Soil conditions, microbial biomass and respiration, mineralization and denitrification rates were measured in upper salt marsh that had been ungrazed or cattle grazed for several decades. Increased microbial biomass and soil respiration were observed in grazed marsh, most likely stimulated by enhanced rates of root turnover and root exudation. We found a significant positive effect of grazing on potential N mineralization rates measured in the laboratory, but this difference did not translate to in situ net mineralization measured monthly from May to September. Rates of denitrification were lowest in the grazed marsh and appeared to be limited by nitrate availability, possibly due to more anoxic conditions and lower rates of nitrification. The major effect of grazing on N cycling therefore appeared to be in limiting losses of N through denitrification, which may lead to enhanced nutrient availability to saltmarsh plants, but a reduced ability of the marsh to act as a buffer for land-derived nutrients to adjacent coastal areas. Additionally, we investigated if grazing influences the rates of turnover of labile and refractory C in saltmarsh soils by adding ¹⁴C-labelled leaf litter or root exudates to soil samples and monitoring the evolution of ¹⁴CO₂. Grazing had little effect on the rates of mineralization of ¹⁴C used as a respiratory substrate, but a larger proportion of ¹⁴C was partitioned into microbial biomass and immobilized in long- and medium-term storage pools in the grazed treatment. Grazing slowed down the turnover of the microbial biomass, which resulted in longer turnover times for both leaf litter and root exudates. Grazing may therefore affect the longevity of C in the soil and alter C storage and utilization pathways in the microbial community.     

缓释复合肥料对土壤和黑麦草氮素营养的影响

采用恒温培养和盆栽生物试验研究了非包膜缓释复合肥料对土壤氮素养分(铵态氮、硝态氮、碱解氮、微生物量氮和固定态铵)和黑麦草氮素营养的效应。结果表明,在恒温培养条件下,各施肥处理土壤铵态氮含量随培养时间均表现为先升高后降低的趋势,且中后期缓释复合肥料处理(SRCF)高于普通复合肥料处理(CCF);缓释复合肥料SRCF1处理土壤硝态氮含量始终低于普通复合肥料CCF1处理,缓释复合肥料SRCF2处理土壤硝态氮含量在初期较高、中后期较低;SRCF各处理土壤碱解氮和微生物量氮含量变化均表现为增加-降低-增加-降低趋势,分别在培养第21d和105d时出现峰值;土壤固定态铵含量变化较小,但总体上以SRCF处理高于CCF处理。在等养分比例盆栽试验中,SRCF1-3处理黑麦草株高、生物量、干重和氮素养分吸收量均高于CCF1处理,SRCF有利于提高黑麦草产量、氮素吸收和利用率;黑麦草氮素吸收量与培养土壤铵态氮、微生物量氮含量呈负相关,而与硝态氮和固定态铵含量呈正相关。     

Isotopic fractionation by saprotrophic microfungi: Effects of species, temperature and the age of colonies

Saprotrophic fungi represent an important resource for a number of fungivorous and omnivorous soil animals, but little is known about the patterns of isotopic fractionation by soil fungi. We grew five common species of saprotrophic microfungi in laboratory cultures on simple artificial substrate based on carbohydrates derived either from C3 or C4 plants. Fungal cultures were kept at 15, 20 or 25 °C. Isotopic composition of carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) in bulk fungal tissue was determined after 11, 21 and 32 days. The fractionation of carbon and nitrogen stable isotopes was species-specific, but generally did not differ in C3- and C4-based growth media. The Zygomycete Mucor plumbeus did not differ in δ¹³C from the carbon source used, though Ascomycetes (Alternaria alternata, Cladosporium cladosporioides, Trichoderma harzianum and Ulocladium botrytis) were depleted in heavy carbon relative to the carbon source by 0.5-0.9‰. Three species were significantly depleted in ¹⁵N relative to the sodium nitrate that was used as a single source of nitrogen. In all species, δ¹⁵N but not δ¹³C tended to increase with the age of fungal colonies. The effect of temperature on δ¹⁵N was weak and inconsistent in different species. In contrast, all fungi except T. harzianum accumulated more ¹³С at 25 °C than at 15 °C. The overall variation in the isotopic signatures of saprotrophic fungi growing in identical conditions reached 8‰ for δ¹⁵N and 2.5‰ for δ¹³C due to species-specific differences in the isotopic fractionation and the age of individual fungal colonies. This variation should be incorporated into the interpretation of the isotopic composition of fungivorous soil animals.