森林凋落物分解及其对全球气候变化的响应

凋落物分解是重要的森林生态系统过程之一,受到气候、凋落物质量、土壤生物群落等生物和非生物因素的综合调控．迄今,有关不同森林生态系统和不同树种地上部分的凋落物动态、凋落物分解过程中的养分释放动态、生物和非生物因素对凋落物分解的影响等研究报道较多,但对地下凋落物的分解研究相对较少．近年来,森林凋落物分解对以大气CO2浓度增加和温度升高为主要特征的全球变化的响应逐步受到重视,但其研究结果仍具有很多不确定性．因此,未来凋落物生态研究的重点应是凋落物分解对土壤有机碳固定的贡献、地上／地下凋落物的物理、化学和生物学过程及其对各种生态因子（例如冻融、干湿交替）及交互作用的响应、凋落物特别是地下凋落物分解对全球气候变化的响应机制等方面．     

Litter mass loss rates in pine forests of Europe and Eastern United States: some relationships with climate and litter quality

The purpose of this study was to relate regional variation in litter mass-loss rates (first year) in pine forests to climate across a large, continental-scale area. The variation in mass-loss rate was analyzed using 39 experimental sites spanning climatic regions from the subarctic to subtropical and Mediterranean: the latitudinal gradient ranged from 31 °N to 70 °N and may represent the the largest geographical area that has ever been sampled and observed for the purpose of studying biogeochemical processes. Because of unified site design and uniform laboratory procedures, data from all sites were directly comparable and permitted a determination of the relative influence of climateversus substrate quality viewed from the perspective of broad regional scales. Simple correlation applied to the entire data set indicated that annual actual evapotranspiration (AET) should be the leading climatic constraint on mass-loss rates (R_adj ² = 0.496). The combination of AET, average July temp. and average annual temp. could explain about 70% of the sites' variability on litter mass-loss. In an analysis of 23 Scots pine sites north of the Alps and Carpatians AET alone could account for about 65% of the variation and the addition of a substrate-quality variable was sufficiently significant to be used in a model. The influence of litter quality was introduced into a model, using data from 11 sites at which litter of different quality had been incubated. These sites are found in Germany, the Netherlands, Sweden and Finland. At any one site most ( ≫ 90%) of the variation in mass-loss rates could be explained by one of the litter-quality variables giving concentration of nitrogen, phosphorus or water solubles. However, even when these models included nitrogen or phosphorus even small changes in potential evapotranspiration resulted in large changes in early-phase decay rates. Further regional subdivision of the data set, resulted in a range of strength in the relationship between loss rate and climatic variables, from very weak in Central Europe to strong for the Scandinavian and Atlantic coast sites (R_adj ² = 0.912; AETversus litter mass loss). Much of the variation in observed loss rates could be related to continentalversus marine/Atlantic influences. Inland locations had mass-loss rates lower than should be expected on the basis of for example AET alone. Attempts to include seasonality variables were not successful. It is clear that either unknown errors and biases, or, unknown variables are causing these regional differences in response to climatic variables. Nevertheless these results show the powerful influence of climate as a control of the broad-scale geography of mass-loss rates and substrate quality at the stand level. Some of these relationships between mass-loss rate and climatic variables are among the highest ever reported, probably because of the care taken to select uniform sites and experimental methods. This suggest that superior, base line maps of predicted mass-loss rates could be produced using climatic data. These models should be useful to predict the changing equilibrium litter dynamics resulting from climatic change.     

Microbial soil respiration and its dependency on carbon inputs, soil temperature and moisture

This experiment was designed to study three determinant factors in decomposition patterns of soil organic matter (SOM): temperature, water and carbon (C) inputs. The study combined field measurements with soil lab incubations and ends with a modelling framework based on the results obtained. Soil respiration was periodically measured at an oak savanna woodland and a ponderosa pine plantation. Intact soils cores were collected at both ecosystems, including soils with most labile C burnt off, soils with some labile C gone and soils with fresh inputs of labile C. Two treatments, dry‐field condition and field capacity, were applied to an incubation that lasted 111 days. Short‐term temperature changes were applied to the soils periodically to quantify temperature responses. This was done to prevent confounding results associated with different pools of C that would result by exposing treatments chronically to different temperature regimes. This paper discusses the role of the above‐defined environmental factors on the variability of soil C dynamics. At the seasonal scale, temperature and water were, respectively, the main limiting factors controlling soil CO₂ efflux for the ponderosa pine and the oak savanna ecosystems. Spatial and seasonal variations in plant activity (root respiration and exudates production) exerted a strong influence over the seasonal and spatial variation of soil metabolic activity. Mean residence times of bulk SOM were significantly lower at the Nitrogen (N)‐rich deciduous savanna than at the N‐limited evergreen dominated pine ecosystem. At shorter time scales (daily), SOM decomposition was controlled primarily by temperature during wet periods and by the combined effect of water and temperature during dry periods. Secondary control was provided by the presence/absence of plant derived C inputs (exudation). Further analyses of SOM decomposition suggest that factors such as changes in the decomposer community, stress‐induced changes in the metabolic activity of decomposers or SOM stabilization patterns remain unresolved, but should also be considered in future SOM decomposition studies. Observations and confounding factors associated with SOM decomposition patterns and its temperature sensitivity are summarized in the modeling framework.     

Global decomposition experiment shows soil animal impacts on decomposition are climate‐dependent

Climate and litter quality are primary drivers of terrestrial decomposition and, based on evidence from multisite experiments at regional and global scales, are universally factored into global decomposition models. In contrast, soil animals are considered key regulators of decomposition at local scales but their role at larger scales is unresolved. Soil animals are consequently excluded from global models of organic mineralization processes. Incomplete assessment of the roles of soil animals stems from the difficulties of manipulating invertebrate animals experimentally across large geographic gradients. This is compounded by deficient or inconsistent taxonomy. We report a global decomposition experiment to assess the importance of soil animals in C mineralization, in which a common grass litter substrate was exposed to natural decomposition in either control or reduced animal treatments across 30 sites distributed from 43°S to 68°N on six continents. Animals in the mesofaunal size range were recovered from the litter by Tullgren extraction and identified to common specifications, mostly at the ordinal level. The design of the trials enabled faunal contribution to be evaluated against abiotic parameters between sites. Soil animals increase decomposition rates in temperate and wet tropical climates, but have neutral effects where temperature or moisture constrain biological activity. Our findings highlight that faunal influences on decomposition are dependent on prevailing climatic conditions. We conclude that (1) inclusion of soil animals will improve the predictive capabilities of region‐ or biome‐scale decomposition models, (2) soil animal influences on decomposition are important at the regional scale when attempting to predict global change scenarios, and (3) the statistical relationship between decomposition rates and climate, at the global scale, is robust against changes in soil faunal abundance and diversity.     

Litter decomposition rates in Canadian forests

The effect of litter quality and climate on the rate of decomposition of plant tissues was examined by the measurement of mass remaining after 3 years’ exposure of 11 litter types placed at 18 forest sites across Canada. Amongst sites, mass remaining was strongly related to mean annual temperature and precipitation and amongst litter types the ratio of Klason lignin to nitrogen in the initial tissue was the most important litter quality variable. When combined into a multiple regression, mean annual temperature, mean annual precipitation and Klason lignin:nitrogen ratio explained 73% of the variance in mass remaining for all sites and tissues. Using three doubled CO₂ GCM climate change scenarios for four Canadian regions, these relationships were used to predict increases in decomposition rate of 4–7% of contemporary rates (based on mass remaining after 3 years), because of increased temperature and precipitation. This increase may be partially offset by evidence that plants growing under elevated atmospheric CO₂ concentrations produce litter with high lignin:nitrogen ratios which slows the rate of decomposition, but this change will be small compared to the increased rate of decomposition derived from climatic changes.     

Regional patterns and controls of leaf decomposition in Australian tropical rainforests

Future climates have the potential to alter decomposition rates in tropical forest with implications for carbon emissions, nutrient cycling and retention of standing litter. However, our ability to predict impacts, particularly for seasonally wet forests in the old world, is limited by a paucity of data, a limited understanding of the relative importance of different aspects of climate and the extent to which decomposition rates are constrained by factors other than climate (e.g. soil, vegetation composition). We used the litterbag method to determine leaf litter decay rates at 18 sites distributed throughout the Australian wet tropics bioregion over a 14‐month period. Specifically, we investigated regional controls on litter decay including climate, soil and litter chemical quality. We used both in situ litter collected from litterfall on site and a standardized control leaf litter substrate. The control litter removed the effect of litter chemical quality and the in situ study quantified decomposition specific to the site. Decomposition was generally slower than for other tropical rainforests globally except in our wet and nutrient‐richer sites. This is most likely attributable to the higher latitude, often highly seasonal rainfall and very poor soils in our system. Decomposition rates were best explained by a combination of climate, soil and litter quality. For in situ litter (native to the site) this included: average leaf wetness in the dry season (LWDS; i.e. moisture condensation) and the initial P content of the leaves, or LWDS and initial C. For control litter (no litter quality effect) this included: rainfall seasonality (% dry season days with 0‐mm rainfall), soil P and mean annual temperature. These results suggest that the impact of climate change on decomposition rates within Australian tropical rainforests will be critically dependent on the trajectory of dry season moisture inputs over the coming decades.     

Keeping up with a warming world; assessing the rate of adaptation to climate change

The pivotal question in the debate on the ecological effects of climate change is whether species will be able to adapt fast enough to keep up with their changing environment. If we establish the maximal rate of adaptation, this will set an upper limit to the rate at which temperatures can increase without loss of biodiversity.The rate of adaptation will primarily be set by the rate of microevolution since (i) phenotypic plasticity alone is not sufficient as reaction norms will no longer be adaptive and hence microevolution on the reaction norm is needed, (ii) learning will be favourable to the individual but cannot be passed on to the next generations, (iii) maternal effects may play a role but, as with other forms of phenotypic plasticity, the response of offspring to the maternal cues will no longer be adaptive in a changing environment, and (iv) adaptation via immigration of individuals with genotypes adapted to warmer environments also involves microevolution as these genotypes are better adapted in terms of temperature, but not in terms of, for instance, photoperiod.Long-term studies on wild populations with individually known animals play an essential role in detecting and understanding the temporal trends in life-history traits, and to estimate the heritability of, and selection pressures on, life-history traits. However, additional measurements on other trophic levels and on the mechanisms underlying phenotypic plasticity are needed to predict the rate of microevolution, especially under changing conditions.Using this knowledge on heritability of, and selection on, life-history traits, in combination with climate scenarios, we will be able to predict the rate of adaptation for different climate scenarios. The final step is to use ecoevolutionary dynamical models to make the link to population viability and from there to biodiversity loss for those scenarios where the rate of adaptation is insufficient.     

Relationship between decomposition of cellulose in the soil and tree stand characteristics in natural boreal forests

The relationship between the decomposition of cellulose placed on and buried in the forest floor and various tree stand characteristics was studied at sites with minimal anthropogenic influence. The 22 study sites, including both forested upland and peatland plots, were clustered in 4 catchments between 61°–69° N in Finland. The stands were 60 to 320 years old and composed of varying proportions of Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies Karst.) and deciduous species (mainly t Betula spp.). Cellulose strips (softwood pulp) were placed on the forest floor surface and buried at four depths down to 5 cm for two 1-year periods and the weight loss measured. Decomposition did not significantly correlate with stand age, but was strongly and positively correlated with stand basal area, mean tree height and stem volume. This was valid at all depths, and even after differences due to climate between catchments had been taken into account. The stem volume of Scots pine dominated plots had the highest correlation. Our results showed that decomposition of organic matter on and in the forest floor is related to the stand characteristics. This relationship should be considered when comparing soil processes different stands, even when comparing stands of the same tree species composition.     

Long-term dynamics of pine and hardwood litter in contrasting environments: toward a global model of decomposition

We analysed data on mass loss after five years of decomposition in the field from both fine root and leaf litters from two highly contrasting trees, Drypetes glauca, a tropical hardwood tree from Puerto Rico, and pine species from North America as part of the Long‐Term Intersite Decomposition Experiment (LIDET). LIDET is a reciprocal litterbag study involving the transplanting of litter from 27 species across 28 sites in North and Central America reflecting a wide variety of natural and managed ecosystems and climates, from Arctic tundra to tropical rainforest. After 5 years, estimated k‐values ranged from 0.032 to 3.734, lengths of Phase I (to 20% mass remaining) from 0.49 to 47.92 years, and fractional mass remaining from 0 to 0.81. Pine litter decomposed more slowly than Drypetes litter, supporting the notion of strong control of substrate quality over decomposition rates. Climate exerted strong and consistent effects on decomposition. Neither mean annual temperature or precipitation alone explained the global pattern of decomposition; variables including both moisture availability and temperature (i.e. actual evapotranspiration and DEFAC from the CENTURY model) were generally more robust than single variables. Across the LIDET range, decomposition of fine roots exhibited a Q₁₀ of 2 and was more predictable than that of leaves, which had a higher Q₁₀ and greater variability. Roots generally decomposed more slowly than leaves, regardless of genus, but the ratio of above‐ to belowground decomposition rates differed sharply across ecosystem types. Finally, Drypetes litter decomposed much more rapidly than pine litter in ‘broadleaved habitats’ than in ‘conifer habitats’, evidence for a ‘home‐field advantage’ for this litter. These results collectively suggest that relatively simple models can predict decomposition based on litter quality and regional climate, but that ecosystem‐specific problems may add complications.     

Interactions between atmospheric CO2 enrichment and soil fauna

We have reviewed the responses of soil fauna to increased concentrations of atmospheric CO₂ and the consequent climate change. These will affect several attributes of animal populations and communities including their density, biomass, diversity, activity, rates of consumption, life history parameters and migration ability. Changes in the quality and quantity of litter and global warming are the main factors which are expected to modify soil fauna. Although changes have been observed in several attributes of the soil fauna as a consequence of increased concentrations of atmospheric CO₂, no general trend which might allow to the prediction of a general pattern of response has been identified. Because of the complexity of the biological mechanisms and the synergetic action of several factors, the few resulting responses reported in the literature are inconclusive. However, some aspects of the situation deserve more attention. These include the consequences of (1) changes in the food resources for soil fauna in the litter layer and in the rhizosphere, (2) the consumption of low quality litter by the macrofauna, (3) the change in life span in response to temperature elevation, (4) the enhancement of earthworm burrowing activity and (5) the changes in community composition arising because of specific differential resistance to adverse conditions. This revised version was published online in June 2006 with corrections to the Cover Date.     

Litter decay controlled by temperature,not soil properties,affecting future soil carbon

Widespread global changes, including rising atmospheric CO₂ concentrations, climate warming and loss of biodiversity, are predicted for this century; all of these will affect terrestrial ecosystem processes like plant litter decomposition. Conversely, increased plant litter decomposition can have potential carbon‐cycle feedbacks on atmospheric CO₂ levels, climate warming and biodiversity. But predicting litter decomposition is difficult because of many interacting factors related to the chemical, physical and biological properties of soil, as well as to climate and agricultural management practices. We applied ¹³C‐labelled plant litter to soil at ten sites spanning a 3500‐km transect across the agricultural regions of Canada and measured its decomposition over five years. Despite large differences in soil type and climatic conditions, we found that the kinetics of litter decomposition were similar once the effect of temperature had been removed, indicating no measurable effect of soil properties. A two‐pool exponential decay model expressing undecomposed carbon simply as a function of thermal time accurately described kinetics of decomposition. (R² = 0.94; RMSE = 0.0508). Soil properties such as texture, cation exchange capacity, pH and moisture, although very different among sites, had minimal discernible influence on decomposition kinetics. Using this kinetic model under different climate change scenarios, we projected that the time required to decompose 50% of the litter (i.e. the labile fractions) would be reduced by 1–4 months, whereas time required to decompose 90% of the litter (including recalcitrant fractions) would be reduced by 1 year in cooler sites to as much as 2 years in warmer sites. These findings confirm quantitatively the sensitivity of litter decomposition to temperature increases and demonstrate how climate change may constrain future soil carbon storage, an effect apparently not influenced by soil properties.     

Daily variations in the optical properties of a small lake

1. The major components of the underwater light field (ULF: vertical attenuation, absorption, scattering and suspensoids including plankton fractions) of Las Madres Lake, a small wind‐sheltered, oligohumic lake in Central Spain, were investigated daily over a period of 3 months at the onset of vernal circulation. 2. Gilvin, arising mostly from the decomposition of reeds in the littoral in autumn, was the main component of vertical attenuation, and its variability explained the highest fraction of absorption variability. Tripton appeared to be the main factor responsible for scattering, and might have resulted from dust deposition from the surrounding mining land. The plankton community played a minor role in attenuation, absorption or scattering throughout the investigation period. 3. Vertical and horizontal mixing dynamics may control the ULF to a certain extent, as most optical properties changed within different mixing periods and poor advective exchanges may have resulted in uneven distribution of water colour in this small lake. 4. Time series analysis showed that most autocorrelations were shorter than a week, inherent properties (absorption, scattering) being delayed longer than apparent properties (attenuation, transparency) as a result of their lower dependence on solar irradiance. A 2‐day lag was observed in cross‐correlations between either gilvin and absorption or tripton and scattering. When different mixing periods at early circulation were considered, however, ULF components changed their relationships and delays with suspensoids and dissolved substances over such periods, probably tracking the dynamics of their controlling factors. 5. Our study, and others at daily, weekly, seasonal, interannual and long‐term scales, demonstrates that ULF is a system upon which different processes are operating at different time scales. Contrary to expectations, however, the variability in the ULF does not increase with time scale and depends partly upon the trophic status of lakes.     

Element interactions in forest ecosystems: succession,allometry and input-output budgets

Element interactions within forests differ from those in other major ecosystems for three major reasons: — a greater allocation of carbon to structural material; — a greater element storage within biomass; and — the diversity of carbon- and nutrient-containing metabolites produced. The most important of these differences is structural material, which can lead to C: element ratios in biomass (as a whole) 100 × greater than those in unicellular organisms. Stand allometry causes the amount of carbon stored and C:element ratios in biomass to change in predictable ways in the course of secondary succession. Such changes affect microbial dynamics and C: element interactions within soils. Bicarbonate, organic acids, nitrate, phosphate, and sulfate are major anions within forest soils: they control leaching of both anions and cations. Biotic interactions of C, N, P, and S during both uptake and mineralization control the potential for production of these anions within forests, and geochemical interactions regulate their mobility and loss.     

Leaf Decay Processes during and after a Supra‐Seasonal Hydrological Drought in a Temperate Lowland Stream

Climate change models for Central Europe predict hydrological drought with fragmentation into pools during periods of high litter input in numerous lowland streams, presumably affecting in‐stream leaf decay processes. To investigate this assumption, we measured physicochemical parameters, macro‐invertebrate colonization, microbial activity, and decay rates of exposed leaves during and after a supra‐seasonal drought in a German lowland stream. Microbial activity, shredder colonization and leaf decay rates during fragmentation were low, presumably caused by drought‐related environmental conditions. Microbial activity and temperature‐corrected decay rates increased after the flow resumption but not leaf mass loss and shredder colonization. During both periods, exposed leaves appeared physically unaffected suggesting strongly reduced shredder‐mediated leaf decay despite shredder presence. Our results indicate that hydrological drought can affect organisms and processes in temperate lowland streams even after flow resumption, and should be considered in climate change scenarios. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

烧失量数值波动对北京地区过去气候和环境的特征响应

通过对北京地区房山东甘池剖面所取的１９７个样品的烧失量测定,结合＾１４Ｃ测年,剖面岩性特征及孢粉序列分析,初步回答了烧失量作为沉积物有机质含量的代用指标,其数值波动对北京地区１５０００ａ以来环境变迁和气候变化的特征响应,同时针对房山东甘池剖面的６个带的烧失量变化特征,提出其在反映气候异常,沉积物地貌单元等的意义,从而为更好重建过去气候和环境提供依据。     

Latitudinal differentiated water table control of carbon dioxide, methane and nitrous oxide fluxes from hydromorphic soils: feedbacks to climate change

The possibility of carbon (C) being locked away from the atmosphere for millennia is given in hydromorphic soils. However, the water-table-dependent feedback from soil organic matter (SOM) decomposition to the climate system is less clear. At least three greenhouse gases are produced: carbon dioxide (CO₂), methane (CH₄) and nitrous oxide (N₂O). These gases show emission peaks at different water table positions and have different global warming potentials (GWP), for example a factor of 23 for CH₄ and 296 for N₂O as compared with the equivalent mass of CO₂ on a 100-year time horizon. This review of available annual data on all three gases revealed that the radiative forcing effect of SOM decomposition is principally dictated by CO₂ despite its low GWP. Anaerobic SOM decomposition generally has a lower potential feedback to the climatic system than aerobic SOM decomposition. Concrete values are constrained by a lack of data from tropical and subarctic regions. Furthermore, data on N₂O and on plant effects are generally rare. However, there is a clear latitudinal differentiation for the GWP of soils under anaerobic conditions compared with aerobic conditions when looking at CO₂ and CH₄: in the tropical and temperate regions, the anaerobic GWP showed a range of 25–60% of the aerobic value, but values varied between 80% and 110% in the boreal zone. Hence, particularly in the vulnerable boreal zone, the feedback from ecosystems to climate change will highly depend on plant responses to changing water tables at elevated temperatures.     

Interactive Effects of Fire, Soil Climate, and Moss on CO2 Fluxes in Black Spruce Ecosystems of Interior Alaska

Fire is an important control on the carbon (C) balance of the boreal forest region. Here, we present findings from two complementary studies that examine how fire modifies soil organic matter properties, and how these modifications influence rates of decomposition and C exchange in black spruce (Picea mariana) ecosystems of interior Alaska. First, we used laboratory incubations to explore soil temperature, moisture, and vegetation effects on CO₂ and DOC production rates in burned and unburned soils from three study regions in interior Alaska. Second, at one of the study regions used in the incubation experiments, we conducted intensive field measurements of net ecosystem exchange (NEE) and ecosystem respiration (ER) across an unreplicated factorial design of burning (2 year post-fire versus unburned sites) and drainage class (upland forest versus peatland sites). Our laboratory study showed that burning reduced the sensitivity of decomposition to increased temperature, most likely by inducing moisture or substrate quality limitations on decomposition rates. Burning also reduced the decomposability of Sphagnum-derived organic matter, increased the hydrophobicity of feather moss-derived organic matter, and increased the ratio of dissolved organic carbon (DOC) to total dissolved nitrogen (TDN) in both the upland and peatland sites. At the ecosystem scale, our field measurements indicate that the surface organic soil was generally wetter in burned than in unburned sites, whereas soil temperature was not different between the burned and unburned sites. Analysis of variance results showed that ER varied with soil drainage class but not by burn status, averaging 0.9 ± 0.1 and 1.4 ± 0.1 g C m⁻² d⁻¹ in the upland and peatland sites, respectively. However, a more complex general linear model showed that ER was controlled by an interaction between soil temperature, moisture, and burn status, and in general was less variable over time in the burned than in the unburned sites. Together, findings from these studies across different spatial scales suggest that although fire can create some soil climate conditions more conducive to rapid decomposition, rates of C release from soils may be constrained following fire by changes in moisture and/or substrate quality that impede rates of decomposition. Author contributions: JAO: performed research, analyzed data, contributed new methods, wrote the paper; MRT: designed laboratory study, performed research, analyzed data; JWH: designed field study, performed research; KLM: performed research; LEP: performed research, contributed new method; GS: performed research; JCN: performed research.     

Mammalian herbivores,grass height and rainfall drive termite activity at different spatial scales in an African savanna

Termites have a large influence on ecosystem functioning. Understanding what drives termite activity patterns improves understanding of nutrient cycling, productivity, and heterogeneity in savannas. We present a mechanistic framework that relates the interactive effects of rainfall, grassland structure, large herbivore presence, and soil factors to termite activity. To test this framework, we used grass litterbags to monitor termite activity at ten sites across Hluhluwe‐iMfolozi Park, South Africa. We assessed the effects of abiotic and biotic factors on termite activity at two scales: the large (landscape) scale, variation in bait removal among 300 m² plots that were distributed across the park and at the small (within‐plot) scale (1–300 m²). Half of our sites were located inside large herbivore exclosures to test for the effect of mammalian herbivore presence. At the landscape scale, termite grass removal declined towards higher rainfall and in the presence of mammalian herbivores. Removal did not depend on soil factors. At the small scale, removal declined with increasing grass height, particularly in the 1 m surrounding the bait bag. Resource quality did not affect bait removal. We suggest that competition for forage drives the negative effect of mammalian herbivores on termites, whereas lower bait removal in taller swards may be due to direct negative effects from rainfall, fire and/or competition with free‐living microbes. Ultimately, we suggest that the impact of termites on nutrient cycling is most pronounced when abiotic (rainfall) and biotic conditions (mammalian herbivory) limit grass removal by fire and decomposition by free‐living microbes.