Delays in nutrient cycling and plant population oscillations

It is well known that delay-differential and delay-difference equations can produce plausible simulations of population oscillations, but many of these equations lack a specific mechanism responsible for the delay. We suggest that delays in release of nitrogen from decomposing litter, caused by microbial uptake, could produce oscillations in populations when the delay in the release of nitrogen is longer than the characteristic time scale of nitrogen uptake. We present a model which captures these dynamics. As the parameter controlling microbial uptake of nitrogen during litter decay increases, the model solutions bifurcate from fixed point equilibria, to periodic orbits (oscillations) which remain bounded for ecologically very long times, and finally to extinction of the plant population because of rapid increases in the amplitude of the oscillations. We suggest that such a mechanism may be especially important for annual plants which do not store nitrogen in perennial tissues to buffer delays. Natural populations of wild rice ( Zizania palustris ), an annual plant, oscillate with approximately four-year periods. Our model qualitatively mimics the period and shape of population oscillations in wild rice with parameter values in the range of those determined by experiments. The model therefore demonstrates a logical and experimentally plausible link between plant population dynamics and the ecosystem processes delaying the cycling of limiting nutrients.     

Carbon and Nitrogen Decoupling Under an 11-Year Drought in the Shortgrass Steppe

The frequency and magnitude of drought is expected to increase in the US Great Plains under future climate regimes. Although semiarid systems are considered highly resistant to water limitation, novel drought events could alter linkages among biogeochemical processes, and result in new feedbacks that influence the timescale of ecosystem recovery. We examined changes in carbon and nitrogen cycling in the last 2 years of an 11-year drought manipulation in the shortgrass steppe, and under the first 2 years of recovery from drought. We measured plant production, plant tissue chemistry, soil trace gas flux, and soil inorganic nitrogen dynamics to test the extent that this magnitude of drought altered carbon and nitrogen fluxes and how these changes affected post-drought dynamics. We found that soil inorganic nitrogen was up to five times higher under severe drought than under control conditions, but that this nitrogen may not have been accessible to plants and microbial communities during drought due to diffusion limitations. Drought plots had higher N₂O flux when they received equal rainfall pulses, showing that this accumulated N may be vulnerable to loss. In addition, plants in drought plots had higher tissue nitrogen for 2 years following drought. These results show that decadal-length droughts that may occur under future precipitation regimes are likely to alter ecosystem properties through interactions among precipitation, vegetation, and N cycling. Shifts in plant N, vulnerability of nitrogen to loss, and rainfall use efficiency that we observed are likely to affect the recovery time of semiarid systems subject to droughts of this magnitude.     

Mechanisms of plant species impacts on ecosystem nitrogen cycling

Plant species are hypothesized to impact ecosystem nitrogen cycling in two distinctly different ways. First, differences in nitrogen use efficiency can lead to positive feedbacks on the rate of nitrogen cycling. Alternatively, plant species can also control the inputs and losses of nitrogen from ecosystems. Our current understanding of litter decomposition shows that most nitrogen present within litter is not released during decomposition but incorporated into soil organic matter. This nitrogen retention is caused by an increase in the relative nitrogen content in decomposing litter and a much lower carbon‐to‐nitrogen ratio of soil organic matter. The long time lag between plant litter formation and the actual release of nitrogen from the litter results in a bottleneck, which prevents feedbacks of plant quality differences on nitrogen cycling. Instead, rates of gross nitrogen mineralization, which are often an order of magnitude higher than net mineralization, indicate that nitrogen cycling within ecosystems is dominated by a microbial nitrogen loop. Nitrogen is released from the soil organic matter and incorporated into microbial biomass. Upon their death, the nitrogen is again incorporated into the soil organic matter. However, this microbial nitrogen loop is driven by plant‐supplied carbon and provides a strong negative feedback through nitrogen cycling on plant productivity. Evidence supporting this hypothesis is strong for temperate grassland ecosystems. For other terrestrial ecosystems, such as forests, tropical and boreal regions, the data are much more limited. Thus, current evidence does not support the view that differences in the efficiency of plant nitrogen use lead to positive feedbacks. In contrast, soil microbes are the dominant factor structuring ecosystem nitrogen cycling. Soil microbes derive nitrogen from the decomposition of soil organic matter, but this microbial activity is driven by recent plant carbon inputs. Changes in plant carbon inputs, resulting from plant species shifts, lead to a negative feedback through microbial nitrogen immobilization. In contrast, there is abundant evidence that plant species impact nitrogen inputs and losses, such as: atmospheric deposition, fire‐induced losses, nitrogen leaching, and nitrogen fixation, which is driven by carbon supply from plants to nitrogen fixers. Additionally, plants can influence the activity and composition of soil microbial communities, which has the potential to lead to differences in nitrification, denitrification and trace nitrogen gas losses. Plant species also impact herbivore behaviour and thereby have the potential to lead to animal‐facilitated movement of nitrogen between ecosystems. Thus, current evidence supports the view that plant species can have large impacts on ecosystem nitrogen cycling. However, species impacts are not caused by differences in plant quantity and quality, but by plant species impacts on nitrogen inputs and losses.     

Biodiversity and decomposition in experimental grassland ecosystems

We examined the impact of biodiversity on litter decomposition in an experiment that manipulated plant species richness. Using biomass originating from the experimental species richness gradient and from a species used as a common substrate, we measured rates of decomposition in litterbags in two locations: in situ in the experiment plots and in an adjacent common garden. This allowed us to separate the effects of litter quality and decomposition location on decomposition. We found that plant species richness had a significant, but minor negative effect on the quality (nitrogen concentration) of the biomass. Neither litter type nor location had a consistent effect on the rate of carbon and nitrogen loss over a 1-year period. Thus, the increased productivity and corresponding lower soil available nitrogen levels observed in high diversity plots do not lead to faster litter decomposition or faster nitrogen turnover. This supports the hypothesis that increased productivity corresponding with higher species richness results in increased litter production, higher standing litter pools and a negative feedback on productivity, because of an increased standing nitrogen pool in the litter.     

The Effects of Exotic Grasses on Litter Decomposition in a Hawaiian Woodland: The Importance of Indirect Effects

Exotic grasses and grass-fueled fires have altered plant species composition in the seasonal submontane woodlands of Hawaii Volcanoes National Park. These changes have altered both structural and functional aspects of the plant community, which could, in turn, have consequences for litter decomposition and nitrogen (N) dynamics. In grass-invaded unburned woodland, grass removal plots within the woodland, and woodland converted to grassland by fire, we compared whole-system fluxes and the contributions of individual species to annual aboveground fine litterfall and litterfall N, and litter mass and net N loss. We assessed the direct contribution of grass biomass to decomposition and N dynamics, and we determined how grasses affected decomposition processes indirectly via effects on native species and alteration of the litter layer microenvironment. Grasses contributed 35% of the total annual aboveground fine litterfall in the invaded woodland. However, total litterfall mass and N were not different between the invaded woodland and the grass removal treatment because of compensation by the native tree Metrosideros polymorpha, which increased litter production by 37% ± 5% when grasses were removed. The 0.3 g N m^–2/y^–1 contained in this production increase was equal to the N contained in grass litter. Litter production and litterfall N was lowest in the grassland due to the loss of native litter inputs. Decomposition of litterfall on an area basis was highest in the grass-invaded woodland. We attributed this effect to increased inherent decomposability of native litter in the presence of grasses because (a) the microenvironment of the three vegetation treatments had little effect on decomposition of common litter types and (b) M. polymorpha litter produced in the invaded woodland decomposed faster than that produced in the grass removal plots due to higher lignin concentrations in the latter than in the former. Area-weighted decomposition was lowest in the grassland due to the absence of native litter inputs. Across all treatments, most litter types immobilized N throughout the incubation, and litter net N loss on an area basis was not different among treatments. Our results support the idea that the effects of a plant species or growth form on decomposition cannot be determined in isolation from the rest of the community or from the direct effects of litter quality and quantity alone. In this dry woodland, exotic grasses significantly altered decomposition processes through indirect effects on the quantity and quality of litter produced by native species.     

Litter Quantity and Nitrogen Immobilization Cause Oscillations in Productivity of Wild Rice (Zizania palustris L.) in Northern Minnesota

Microbial immobilization of nitrogen (N) in litter from one year’s production may cause oscillations in biomass production if it delays N availability the following year. We tested whether shoot and root litter and plant density affect biomass and seed production of populations of wild rice (Zizannia palustris L.) grown in 378 l stock tank mesocosms over four consecutive years. Half the tanks were thinned to a uniform seedling density whereas density in the remaining tanks was allowed to fluctuate ad libitum. Litter treatments included both shoot litter removal, leaving only root litter, and retaining shoot litter intact with root litter. A separate greenhouse fertilizer experiment tested whether N and/or phosphorus (P) limited productivity. Responses to N additions were much stronger than to P additions. Annual production and N availability in the tanks were correlated with each other and followed a concurrent cycle from 2004 to 2008. Furthermore, production in tanks with shoot + root litter did not fluctuate more than tanks with only root litter. Root litter immobilized more nitrogen and for a longer period than shoot litter. Neither litter immobilized P. Density did not affect mean seed weight, total seed production, or mean plant weight, but total seed production declined in years following productive years and was high only following years of low litter production. Root litter may therefore be primarily responsible for the delays in N availability that cause cycles in biomass and seed production. Consequently, both wild rice litter quantity and quality play central roles in production and population dynamics of wild rice stands.     

Disentangling the Litter Quality and Soil Microbial Contribution to Leaf and Fine Root Litter Decomposition Responses to Reduced Rainfall

Climate change-induced rainfall reductions in Mediterranean forests negatively affect the decomposition of plant litter through decreased soil moisture. However, the indirect effects of reduced precipitation on litter decomposition through changes in litter quality and soil microbial communities are poorly studied. This is especially the case for fine root litter, which contributes importantly to forests plant biomass. Here we analyzed the effects of long-term (11 years) rainfall exclusion (29% reduction) on leaf and fine root litter quality, soil microbial biomass, and microbial community-level physiological profiles in a Mediterranean holm oak forest. Additionally, we reciprocally transplanted soils and litter among the control and reduced rainfall treatments in the laboratory, and analyzed litter decomposition and its responses to a simulated extreme drought event. The decreased soil microbial biomass and altered physiological profiles with reduced rainfall promoted lower fine root—but not leaf—litter decomposition. Both leaf and root litter, from the reduced rainfall treatment, decomposed faster than those from the control treatment. The impact of the extreme drought event on fine root litter decomposition was higher in soils from the control treatment compared to soils subjected to long-term rainfall exclusion. Our results suggest contrasting mechanisms driving drought indirect effects on above-(for example, changes in litter quality) and belowground (for example, shifts in soil microbial community) litter decomposition, even within a single tree species. Quantifying the contribution of these mechanisms relative to the direct soil moisture-effect is critical for an accurate integration of litter decomposition into ecosystem carbon dynamics in Mediterranean forests under climate change.     

Invasive grass litter facilitates native shrubs through abiotic effects

Questions: Plant invasions are considered one of the top threats to the biodiversity of native taxa, but clearly documenting the causal links between invasions and the decline of native species remains a major challenge of invasion biology. Most studies have focused on impacts of invaders' living biomass, rather than on mechanisms mediated by litter. However, invasive plant litter, which is often of a very different type and quantity than a system's native plant litter, can have multiple important effects on ecosystem processes – such as nitrogen cycling and soil microclimate – that may influence native plants. Location: We studied effects of litter of invasive grass species that are widespread throughout western North America on native shrubs in southern California's semi‐arid habitat of coastal sage scrub. Methods: We combined a 3‐year field manipulation of non‐native litter with structural equation modeling to understand interacting effects on non‐native grasses, native shrubs, soil nitrogen (available and total), and soil moisture. Results: Litter addition facilitated non‐native grass growth, revealing a positive feedback likely to enhance invasion success. Contrary to a major paradigm of invasion biology – that competition with invasive plant species causes declines of native plants – we found that litter also facilitated growth of the native dominant shrub, a result supported by observational trends. Structural equation models indicated that enhanced soil moisture mediated the positive effects of litter on shrub growth. Conclusions: We demonstrate that invasive plants, via their litter, can facilitate dominant native plants by altering soil moisture. Our results highlight that understanding the impacts and mechanisms of plant invasions may be enhanced by considering the role of invasive plant litter on native plants and ecosystem properties.     

Plant–fungal symbiosis affects litter decomposition during primary succession

Microbial symbionts of plants can affect decomposition by altering the quality or quantity of host plant tissue (substrate) or the micro‐environment where decomposition occurs (conditioning). In C₃ grasses, foliar fungal endophytes (Clavicipitaceae) can increase plant resistance to drought and/or produce alkaloids that reduce herbivory – effects that may also influence host litter composition and subsequent litter decomposition. We studied the effect of the endophyte Epichloë sp. on litter decomposition in the Great Lakes dunes (USA) using a reciprocal design altering endophyte presence/absence in both American beachgrass Ammophila breviligulata substrate (litter bags) and its conditioning of the decomposition microenvironment. Symbiont treatments were crossed with rain‐out shelters that altered growing season precipitation. The first year of decomposition, senesced leaf substrate from A. breviligulata with Epichloë decomposed 21% faster than endophyte‐free substrate. By the third year, conditioning by live symbiotic plants reduced cumulative decomposition by 33% compared to plots planted with endophyte‐free plants. Of the traits we examined – litter quantity, C:N ratio, mineral composition, fungal colonization, and carbon chemistry – increased litter quantity via greater tiller production was the primary trait shift associated with endophyte symbiosis. Epichloë in A. breviligulata litter also altered litter nitrogen decomposition dynamics, as evidenced by lower nitrogen and protein content in decomposed tissue from plants that hosted the endophyte. Differences in initial litter quality and subsequent colonization by saprotrophic fungi were ruled out as key drivers. Altered precipitation had negligible effects on decomposing processes in the dunes. Grass–Epichloë symbiosis altered nutrient cycling through increasing the rate of litter decomposition when present in the litter and through reducing litter decomposition by conditioning the decomposition microenvironment. Epichloë are widespread symbionts of grasses. Thus, their effects on decomposition could be an important, but often overlooked, driver of nutrient cycling in grass‐dominated ecosystems.     

Indirect effects of deer herbivory on local nitrogen availability in a coastal dune ecosystem

Herbivores can have indirect effects on local nutrient availability if their direct effects on plants lead to changes in the amount or chemical composition of litter reaching the soil surface. Using two exclosure experiments, we evaluated this possibility for black-tailed deer ( Odocoileus hemionus columbianus ) feeding on silver bush lupine ( Lupinus chamissonis ) in a coastal dune system in northern California. Our first experiment assessed the effects of deer herbivory on 360 lupines that were tracked from newly germinated seedlings in 1996 and 1997 until 2000, when most surviving individuals had reached reproductive age. Results from this experiment showed that browsing by deer significantly reduced growth rates and seed production of lupines and delayed the onset of reproductive maturity. Although deer had no effect on the density of litter accumulating underneath shrubs (g m⁻²), browsing significantly decreased C:N ratios of leaf tissue, due primarily to increased nitrogen content of leaves. Deer browsing also caused significantly increased net nitrogen mineralization rates in the soil under shrubs, although pools of ammonium and nitrate were uninfluenced. The second experiment examined 72 established bush lupines from 1997 to 2000, and showed that deer browsing significantly decreased seed production but had no effect on shrub growth. We also detected trends for browsed shrubs to have reduced pools of ammonium and nitrate underneath their canopies, although no such patterns emerged for nitrogen mineralization rates. Collectively, these data suggest that changes in litter quality, more so than litter quantity, contributed to the indirect effects of deer herbivory on local nitrogen dynamics. We hypothesize that deer browsing induced chemical changes in lupine plant tissue, which increased the nitrogen content of litter reaching the soil surface and subsequently increased rates of nitrogen mineralization.     

不同土壤厚度、水分和种植方式对喀斯特两种草本凋落物分解质量损失和化学计量特征的影响

为了探究生长期间不同土壤厚度、水分及种植方式处理对草本植物凋落物分解质量损失和化学计量特征的的影响,采用分解袋法,在露天分解床上分解经过生长期间2种土壤厚度(对照土壤厚度和浅土处理)、2种水分(正常灌水处理和干旱处理)和2种种植方式(单种和混种)处理的苇状羊茅(Festuca arundinacea Schreb.)和黑麦草(Lolium perenne L.)凋落物,研究生长期间土壤厚度和水分减少及不同的种植方式是否通过改变两物种初始凋落物质量、产量和组分来影响自身凋落物分解。结果发现:(1)与对照组(CK)相比,在干旱组(D)和浅土+干旱组(SD),两物种地上、根系和总的凋落物质量损失率、初始N和P含量均显著增加,凋落物产量、C/N和C/P显著降低,凋落物地上组分比大体上无显著变化;而不同的种植方式处理对各组分质量损失率、元素含量、计量比、凋落物产量和组分比大体上无显著影响;(2)两物种地上、根系和总凋落物的质量损失率分别与地上、根系和总N含量呈显著正相关,与C/N呈显著负相关,与凋落量呈显著负相关,而总凋落物质量损失率与地上凋落物组分比呈显著正相关。结果表明,生长期间干旱和浅土+干旱处理能够通过影响苇状羊茅和黑麦草的初始凋落物质量、产量和组分比来加快地上、根系和总凋落物分解,其中凋落物N含量和C/N是影响两物种凋落物分解快慢的主要原因。     

Selective gopher disturbance influences plant species effects on nitrogen cycling

Plant species effects on ecosystem processes are mediated by traits such as litter quality and exudation. These same traits also influence the activity and distribution of animals that play key roles in regulating ecosystem dynamics. We planted monocultures of eight plant species commonly found in California grasslands to investigate the relative importance of plant species direct effects on nitrogen cycling, versus their indirect effects mediated by plant interactions with gophers. Plant species differed in their litter C:N ratio, which closely related to species effects on rates of net mineralization and nitrification in undisturbed soil. However, the effect of selective gopher disturbance on N cycling greatly altered these species effects.
Plant species differed in their effects on the type and timing of gopher disturbance. Small feeding holes were formed in late spring in plots containing species with high tissue quality. These feeding holes minimally disturbed the soil and did not alter N cycling rates over the short term. Large gopher mounds were formed in the winter and early spring, primarily in plots containing the grass, Aegilops triuncialis , and to a lesser extent in plots containing Avena barbata . These large mounds significantly disturbed the soil and greatly increased net nitrification rates, but had no consistent effects on net N mineralization. In undisturbed soil, Aegilops had the highest litter C:N ratio and one of the lowest rates of net nitrification. However, gophers preferentially built large mounds in Aegilops plots. Once the effects of gopher burrowing were considered, Aegilops had one of the highest rates of net nitrification, indicating that the indirect effects of plant species on N cycling can be more important than the direct effects alone. This experiment indicates that it is vital to consider interactions between plants and other organisms in order to predict the ecosystem effects of plant communities.     

Does light exposure make plant litter more degradable?

Many field experiments have indicated that litter decomposition in semi-arid areas may be partly or fully controlled by photodegradation. We devised a study to test our hypothesis that light exposure makes plant litter more degradable. Dry, senescent, aboveground plant litter from Miscanthus x giganteus was exposed to light including ultraviolet (UV) radiation for various lengths of time from 0 to 289 days. Weight loss was measured after exposure and appeared to be modest and did not increase with time of exposure. The litter of the longest and shortest exposure time as well as controls were then incubated with soil and moisture for 35 days and CO₂ and N₂O production were measured. The longest exposed litter degraded much faster than any other treatment during incubation with moisture, about twice as fast as the unexposed control. The shortest exposed however, degraded only slightly faster than the unexposed control. This suggests that increasing litter degradability is a more important mechanism for photodegradation than direct light-induced mass loss. N₂O production from decomposition of the exposed litter was high in the beginning, suggesting that nitrogen may be released quickly. The mechanism is probably that light exposure leaves the nitrogen in plant litter easily available to microbial utilisation upon wetting. Such a mechanism might play an important role for nutrient cycling in semi-arid areas.     

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.     

Nonnative plant shifts functional groups of arthropods following drought

Nonnative plants alter the composition of native plant communities, with concomitant effects on arthropods. However, plant invasions may not be the only disturbance affecting native communities, and multiple disturbances can have compounding effects. We assessed the effects of invasion and drought on plant and arthropod communities by comparing grasslands dominated by nonnative Old World bluestem grasses (OWBs, Dichanthium annulatum) to grasslands dominated by native plants during a period of decreasing drought severity (2011–2013). Native plant communities had more species of plants and arthropods (/m²) than areas dominated by OWBs during extreme drought, but richness was comparable as drought severity decreased. Abundance of arthropods was greater in native plant communities than in OWB communities during extreme drought, but OWB communities had more arthropods during moderate and non-drought conditions. We observed a shift in the arthropod community from one dominated by detritivores to one dominated by herbivores following plant invasion; the magnitude of this shift increased as drought severity decreased. Both plant communities were dominated by nonnative arthropods. A nonnative leafhopper (Balclutha rubrostriata) and native mites (Mochlozetidae) dominated OWB communities as drought severity decreased, and OWBs may serve as refugia for both taxa. Nonnative woodlice (Armadillidium vulgare) dominated native plant communities during extreme and non-drought conditions and abundance of this species may be associated with an increase in plant litter and available nutrients. Given the importance of arthropods for ecosystem services, incorporating arthropod data into conservation studies may demonstrate how changes in arthropod diversity alter ecosystem function where nonnative plants are dominant.     

Nitrogen losses from perennial grass species

Nitrogen losses from plants may occur through a variety of pathways, but so far, most studies have only quantified losses of nutrients by above-ground litter production. We used ¹⁵N pulse labelling to quantify total nitrogen losses from above- and below-ground plant parts. Using this method we were able to include also pathways other than above-ground litter production. To test the hypothesis that species from nutrient-poor habitats lose less nitrogen than species from more fertile soils, six perennial grasses from habitats with a wide range of nutrient availability were investigated: Lolium perenne, Arrhenatherum elatius, Anthoxanthum odoratum, Festuca rubra, F. ovina and Molinia caerulea. The results of an experiment carried out in pots in a green-house at two fertility levels show that statistically significant losses occur through pathways other than above-ground litter production. In the low fertility treatment, most (70%) losses from L. perenne occurred by litter production, but in Ar. elatius, F. rubra, F. ovina and M. caerulea, more than 50% of labelled N losses took place by root turn-over, leaching or exudation from roots. When nutrient supply increased, the ¹⁵N losses in above-ground dead material increased in all species and in Ar. elatius, A. odoratum and F. rubra the ¹⁵N losses via other pathways decreased. Ranked according to decreasing turnover coefficient the sequence of species was: L. perenne, A. odoratum, F. rubra, F. ovina, Ar. elatius, M. caerulea. These results suggest that species adapted to sites with low availability of nutrients lose less nitrogen (including above- and below-ground losses) than species adapted to more fertile soils.     

Identification of General Patterns of Nutrient and Labile Carbon Control on Soil Carbon Dynamics Across a Successional Gradient

Carbon (C) inputs and nutrient availability are known to affect soil organic carbon (SOC) stocks. However, general rules regarding the operation of these factors across a range of soil nutrient availabilities and substrate qualities are unidentified. “Priming” (stimulated decomposition by labile C inputs) and ‘preferential substrate utilization’ (retarded decomposition due to shifts in community composition towards microbes that do not mineralize SOC) are two hypotheses to explain effects of labile C additions on SOC dynamics. For effects of nutrient additions (nitrogen and phosphorus) on SOC dynamics, the stoichiometric (faster decomposition of materials of low carbon-to-nutrient ratios) and ‘microbial mining’ (that is, reduced breakdown of recalcitrant C forms for nutrients under fertile conditions) hypotheses have been proposed. Using the natural gradient of soil nutrient availability and substrate quality of a chronosequence, combined with labile C and nutrient amendments, we explored the support for these contrasting hypotheses. Additions of labile C, nitrogen (N), phosphorus (P), and combinations of C and N and C and P were applied to three sites: 2-year fallow grassland, mature grassland and forest, and the effects of site and nutrient additions on litter decomposition and soil C dynamics were assessed. The response to C addition supported the preferential substrate hypothesis for easily degradable litter C and the priming hypothesis for SOC, but only in nitrogen-enriched soils of the forest site. Responses to N addition supported the microbial mining hypothesis irrespective of C substrate (litter or SOC), but only in the forest site. Further, P addition effects on SOC support the stoichiometric hypothesis; P availability appeared key to soil C release (priming) in the forest site if labile C and N is available. These results clearly link previously contrasting hypotheses of the factors controlling SOC with the natural gradient in litter quality and nutrient availability that exists in ecosystems at different successional stages. A holistic theory that incorporates this variability of responses, due to different mechanisms, depending on nutrient availability and substrate quality is essential for devising management strategies to safeguard soil C stocks.     

Functional dissimilarity across trophic levels as a driver of soil processes in a Mediterranean decomposer system exposed to two moisture levels

The role of biodiversity for soil processes remains poorly understood. Existing evidence suggests that functional diversity rather than species richness is relevant for soil functioning. However, the importance of functional diversity has rarely been assessed simultaneously at more than one trophic level, critically limiting the prediction of consequences of biodiversity loss for soil functioning. In a laboratory microcosm experiment, we tested the hypothesis that increasing functional dissimilarity of both litter‐feeding soil fauna and litter mixtures interactively affects the rates of five different soil processes related to litter decomposition. We created trait‐based functional dissimilarity gradients using five assemblages of two detritivore species and five mixtures of two plant litter species commonly found in Mediterranean shrubland ecosystems of southern France. With increasing drought periods predicted for Mediterranean ecosystems in the future, we additionally included two different watering frequencies to evaluate the impact of drought on soil processes and how drought interacts with functional dissimilarity. The different fauna assemblages and litter mixtures showed strong effects on litter mass loss, soil organic carbon and nitrogen leaching, as well as on soil microbial activities. Up to 20% of the variation in response variables was explained by functional dissimilarity, suggesting an ecologically relevant impact of functional diversity on soil process rates. Detritivore functional dissimilarity tended to have stronger effects when combined with increasingly dissimilar litter mixtures, suggesting that trait dissimilarity interacts across trophic levels. Drought affected several soil processes but did not modify the relationships between functional dissimilarity and process rates. Our results indicate that trait diversity of detritivore assemblages and litter mixtures is an important predictor of soil process rates. The common and easily measurable traits used in our study suggest straightforward application across different types of ecosystems and environmental conditions.     

The effects of drought and herbivory on plant–herbivore interactions across 16 soybean genotypes in a field experiment

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N2-Fixing Red Alder Indirectly Accelerates Ecosystem Nitrogen Cycling

Symbiotic N₂-fixing tree species can accelerate ecosystem N dynamics through decomposition feedbacks via both direct and indirect pathways. Direct pathways include the production of readily decomposed leaf litter and increased N supply to decomposers, whereas indirect pathways include increased tissue N and altered detrital dynamics of non-fixing vegetation. To evaluate the relative importance of direct and indirect pathways, we compared 3-year decomposition and N dynamics of N₂-fixing red alder leaf litter (2.34% N) to both low-N (0.68% N) and high-N (1.21% N) litter of non-fixing Douglas-fir, and decomposed each litter source in four forests dominated by either red alder or Douglas-fir. We also used experimental N fertilization of decomposition plots to assess elevated N availability as a potential mechanism of N₂-fixer effects on litter mass loss and N dynamics. Direct effects of N₂-fixing red alder on decomposition occurred primarily as faster N release from red alder than Douglas-fir litter. Direct increases in N supply to decomposers via experimental N fertilization did not stimulate decomposition of either species litter. Fixed N indirectly influenced detrital dynamics by increasing Douglas-fir tissue and litter N concentrations, which accelerated litter N release without accelerating mass loss. By increasing soil N, tissue N, and the rate of N release from litter of non-fixers, we conclude that N₂-fixing vegetation can indirectly foster plant–soil feedbacks that contribute to the persistence of elevated N availability in terrestrial ecosystems.