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Biological decomposition and wildfire are two predominant and alternative processes that can mineralize organic C in forest litter. Currently, the relationships between decomposition and fire are still poorly understood.We provide an empirical test of the hypothesized decoupling of surface litter bed decomposability and flammability, and the underlying traits and trait spectra.We employed a 41‐species set of gymnosperms of very broad evolutionary and geographic spread, because of the wide range of (absent to frequent) fire regimes they are associated with.We found that the interspecific pattern of mass loss proportions in a "common garden" decomposition experiment was not correlated with any of the flammability parameters and an RDA analysis also showed that the decomposability and flammability of leaf litter in litter layers were decoupled across species. This decoupling originates from the former depending mostly on size and shape spectrum traits and the latter on PES traits and those trait spectra being virtually uncorrelated.Synthesis: Our results show that, indeed, leaf litter decomposability and flammability parameters are decoupled across species, and this decoupling can be explained by their different drivers in terms of trait spectra: chemical traits for decomposability and size‐shape traits for litter layer flammability. [ABSTRACT FROM AUTHOR]

1 The decomposition rate of litter mixtures can depend on interactions between compounds in the component species, i.e. in litter chemistry. Differences in litter N concentration are likely to lead to positive interaction (i.e. faster than expected decomposition) and differences in phenolic concentrations to a negative interaction. These interactions may become less positive (or more negative) when the litter is produced under elevated atmospheric C[O.sub.2] concentrations. 2 We measured respiration rates of litter from two contrasting species pairs, produced under ambient and elevated C[O.sub.2]. 3 As expected, mixtures of Calamagrostis epigejos and Vicia lathyroides, which differed strongly in litter N concentration, decomposed faster than expected, but the interaction disappeared under elevated C[O.sub.2]. 4 Despite the large difference in litter N concentration between Sphagnum recurvum and Carex rostrata, no interaction was observed in the ambient C[O.sub.2] litter mixtures, and under elevated C[O.sub.2], there was an unexpected positive interaction. 5 Interactions in litter mixtures can therefore change under elevated C[O.sub.2], albeit not in a predictable manner, which would require correct prediction of species-specific litter chemistry and its response to elevated C[O.sub.2]. Key-words: global change, litter decomposition, nutrient cycling

Contents653I.654II.657III.659IV.661V.662VI.663VII.665665References665 Summary: Biological decomposition and wildfire are connected carbon release pathways for dead plant material: slower litter decomposition leads to fuel accumulation. Are decomposition and surface fires also connected through plant community composition, via the species’ traits? Our central concept involves two axes of trait variation related to decomposition and fire. The ‘plant economics spectrum’ (PES) links biochemistry traits to the litter decomposability of different fine organs. The ‘size and shape spectrum’ (SSS) includes litter particle size and shape and their consequent effect on fuel bed structure, ventilation and flammability. Our literature synthesis revealed that PES ‐ driven decomposability is largely decoupled from predominantly SSS ‐ driven surface litter flammability across species; this finding needs empirical testing in various environmental settings. Under certain conditions, carbon release will be dominated by decomposition, while under other conditions litter fuel will accumulate and fire may dominate carbon release. Ecosystem ‐ level feedbacks between decomposition and fire, for example via litter amounts, litter decomposition stage, community ‐ level biotic interactions and altered environment, will influence the trait ‐ driven effects on decomposition and fire. Yet, our conceptual framework, explicitly comparing the effects of two plant trait spectra on litter decomposition vs fire, provides a promising new research direction for better understanding and predicting Earth surface carbon dynamics. [ABSTRACT FROM AUTHOR]

It has long been recognized that leaf traits exert a crucial control on litter decomposition, a key process for nutrient cycling, and that invading species can greatly alter such soil processes via changes in mixed litter trait composition. Trait effects on ecosystem processes are hypothesized to operate via changes in either dominant trait values in the community (often calculated as community-weighted mean trait values; CWM) or trait functional diversity (dissimilarity between species trait values; FD). Few have studied the effects of these community trait components in tandem due to their interdependence., We studied litter mixture decomposition using three exotic and six native European tree species with a range in litter decomposability, to disentangle the unique and combined roles of CWM and FD in explaining net litter mixture mass loss., We showed that while CWM exerted the strongest effect on mass loss, FD modulated its effects, increasing mass loss in mixtures with low mean decomposability and decreasing mass loss in mixtures with high mean decomposability. Litter species identity and native/exotic status explained relatively little additional variation in mass loss after accounting for CWM and FD. We further showed that alterations to CWM and FD were more important than the replacement of a native species with an exotic counterpart in predicting mass loss., Synthesis: Our results indicate that the effect of adding an exotic or losing a native species on litter decomposition rate can be predicted from how a species alters both CWM and FD trait values. This supports the idea that the repercussions of exotic species on ecosystem processes depends on the extent that introduced species bear novel traits or trait values and so on how functionally dissimilar a species is compared to the existing species in the community. [ABSTRACT FROM AUTHOR]

The decomposition of dead organic matter is a major determinant of carbon and nutrient cycling in ecosystems, and of carbon fluxes between the biosphere and the atmosphere. Decomposition is driven by a vast diversity of organisms that are structured in complex food webs. Identifying the mechanisms underlying the effects of biodiversity on decomposition is critical given the rapid loss of species worldwide and the effects of this loss on human well-being. Yet despite comprehensive syntheses of studies on how biodiversity affects litter decomposition, key questions remain, including when, where and how biodiversity has a role and whether general patterns and mechanisms occur across ecosystems and different functional types of organism. Here, in field experiments across five terrestrial and aquatic locations, ranging from the subarctic to the tropics, we show that reducing the functional diversity of decomposer organisms and plant litter types slowed the cycling of litter carbon and nitrogen. Moreover, we found evidence of nitrogen transfer from the litter of nitrogen-fixing plants to that of rapidly decomposing plants, but not between other plant functional types, highlighting that specific interactions in litter mixtures control carbon and nitrogen cycling during decomposition. The emergence of this general mechanism and the coherence of patterns across contrasting terrestrial and aquatic ecosystems suggest that biodiversity loss has consistent consequences for litter decomposition and the cycling of major elements on broad spatial scales. [ABSTRACT FROM AUTHOR]

There is substantial evidence that the rate of litter decomposition is affected by the match between the litter substrate and the soil matrix (decomposer community). We introduce and test the phenology-substrate-match hypothesis, which predicts that both litter composition and soil matrix will change over the course of the year and that a lagged match between litter type and soil matrix will result in an optimal decomposition environment., We conducted a decomposition experiment in a Mediterranean mixed deciduous-evergreen oak savanna in California. We initiated litter decomposition of both a deciduous oak (whose leaves fall in autumn) and an evergreen oak (whose leaves fall in spring) in both autumn and spring., Consistent with the phenology-substrate-match hypothesis, we found that decomposition of deciduous oak litter was accelerated compared to evergreen oak litter when decomposition was initiated in spring, while evergreen litter was accelerated compared to deciduous litter when decomposition was initiated in autumn., We also found a small effect of microsite on leaf decomposition, where both evergreen and deciduous oak leaves decomposed faster under the canopy of a conspecific., Synthesis. Our study extends theory of litter quality and the decomposer community into a temporal context, which may be an important source of variation in decomposition rates when species with different litterfall phenologies co-occur. [ABSTRACT FROM AUTHOR]

The decomposition rates of plant litter mixtures often deviate from the averaged rates of monocultures of their component litter species. The mechanisms behind these non-additive effects in decomposition of litter mixtures are lively debated. One plausible explanation for non-additive effects is given by the improved microenvironmental condition (IMC) theory. According to this theory, plant litter species, whose physical characteristics improve the microclimatic conditions for decomposers, will promote the decomposition of their co-occurring litter species. We tested the IMC theory in relation to leaf litter and soil moisture in two contrasting moisture conditions in a dry subarctic mountain birch forest with vascular plant leaf litters of poor and high quality. The non-additive effects in mass loss of litter mixtures increased when moisture conditions in litter and soil became more favourable for plant litter decomposition. The sign of this increase (antagonistic or synergistic) in non-additive effects was more predictable for litter mixtures of poor litter quality. Although the specific mechanisms underlying the IMC theory depended on the litter quality of the litter mixtures, a standardized water holding capacity (WHC) was the litter trait most closely related to the non-additive effects in mixtures of both poor and high quality litter types. Furthermore, we found that higher dissimilarity in WHC traits between the component litter species in a mixture increased synergistic effects in litter mixtures under limiting moisture conditions. However, under improved moisture conditions, increased antagonistic effects were observed. Thus, we found clear support for the IMC theory and showed that climatic conditions and leaf litter physical traits determine whether the non-additive effects in litter mixtures are antagonistic or synergistic. Our study emphasizes the need to include litter physical traits into predictive models of mixing effects on plant litter decomposition and in general suggests climate specificity into these models. [ABSTRACT FROM AUTHOR]

Plant litter decomposition is a key process in terrestrial carbon cycling, yet the relative importance of various control factors remains ambiguous at a global scale. A full reciprocal litter transplant study with 16 litter species that varied widely in traits and originated from four forest sites covering a large latitudinal gradient (subarctic to tropics) showed a consistent interspecific ranking of decomposition rates. At a global scale, variation in decomposition was driven by a small subset of litter traits (water saturation capacity and concentrations of magnesium and condensed tannins). These consistent findings, that were largely independent of the varying local decomposer communities, suggest that decomposer communities show little specialisation and high metabolic flexibility in processing plant litter, irrespective of litter origin. Our results provide strong support for using trait-based approaches in modelling the global decomposition component of biosphere-atmosphere carbon fluxes. [ABSTRACT FROM AUTHOR]

Summary 1. Recent evidence indicates tight control of plant resource economics over interspecific trait variation amongst species, both within and across organs, referred to as 'plant economics spectrum' (PES). Whether and how these coordinated whole-plant economics strategies can influence the decomposition system and thereby impact on ecosystem carbon and nutrient cycling are yet an open question. More specifically, it is yet unknown whether plant functional traits have consistent afterlife effects across different plant organs. 2. To answer those questions, we conducted a common-garden decomposition experiment bringing together leaves, fine stems, coarse stems, fine roots and reproductive parts from a wide range of subarctic plant types, clades and environments. We measured all plant parts for the same (green and litter) plant economics traits and identified a whole-plant axis of carbon and nutrient economics. 3. We demonstrated that our local 'PES' has important afterlife effects on carbon turnover by driving coordinated decomposition rates of different organs across species. All organ decomposabilities were consistently controlled by the same structure-related traits (lignin, C and dry matter content) whilst nutrient-related traits (N, P, pH, phenols) had more variable influence, likely due to their contrasting functions across organs. Nevertheless, consistent shifts in elevation of parallel trait-decomposition relationships between organs indicate that other variables, potentially related to organ dimensions, configuration or chemical contents, codetermine litter decomposition rates. 4. Whilst the coordinated litter decomposabilities across species organs imply a coordinated impact of plant above-ground and below-ground litters on plant-soil feedbacks, the contrasting decomposabilities between plant parts suggest a major role for the relative inputs of organ litter as driver of soil properties and ecosystem biogeochemistry. These relationships, underpinning the afterlife effects of the PES on whole-plant litter decomposability, will provide comprehensive input of vegetation composition feedback to soil carbon turnover. [ABSTRACT FROM AUTHOR]

Summary 1. While the importance of wood decay for the global carbon balance is widely recognized, surprisingly little is known about its long-term dynamics and its abiotic and biotic drivers. Progress in this field is hindered by the long time-scales inherent to the low decay rates of wood and the lack of short-term methods to assess long-term decomposition dynamics in standardized field conditions. 2. Here, we present such a method, which relies on the sampling and short-term incubation of wood from several decay stages covering the entire decay process. Together these short-term decay steps are used to model and discriminate between three potential decay dynamics (linear, exponential and sigmoid) using an iterative optimization procedure. We applied this method to analyse long-term wood decay of six subarctic tree species (six stems and two roots) and test the hypotheses that (i) different wood species follow distinct decay dynamics and (ii) interspecific variation in wood traits controls variation in wood decay rates in a standardized environment. 3. We found interspecific variation in long-term wood decay dynamics: decay of Alnus and Salix stems was best described by exponential models, whereas decay of Sorbus stems and Betula and Pinus roots was best fitted by linear models and Betula, Pinus and Populus stems each displayed a sigmoid decay dynamics (up to 5-year initial lag phase). A six-fold variation was observed between the decomposition half-lives of all eight wood types, from 6.8 years (6.1-7.5, 95% C.I.) for Alnus stems to 41.3 years (34.5-51.8) for Pinus roots. Initial wood traits such as pH ( R2 = 0.92), dry matter content ( R2 = 0.79) and lignin ( R2 = 0.73) were good predictors of long-term wood decay rates. 4. Synthesis. Our findings suggest changing decay dynamics across wood species and types that are likely to arise from changing underlying wood decay processes (i.e. varying wood functional traits/decomposer community interactions). Our new method, which combines advantages of direct observations and the chronosequence approach, allows reliable comparisons of species contributions to long-term wood decay rates and provides future opportunities to experimentally disentangle intrinsic and external abiotic and biotic drivers of long-term wood decay processes. [ABSTRACT FROM AUTHOR]

As a result of low decomposition rates, high-latitude ecosystems store large amounts of carbon. Litter decomposition in these ecosystems is constrained by harsh abiotic conditions, but also by the absence of macro-detritivores. We have studied the potential effects of their climate change-driven northward range expansion on the decomposition of two contrasting subarctic litter types. Litter of Alnus incana and Betula pubescens was incubated in microcosms together with monocultures and all possible combinations of three functionally different macro-detritivores (the earthworm Lumbricus rubellus, isopod Oniscus asellus, and millipede Julus scandinavius). Our results show that these macro-detritivores stimulated decomposition, especially of the high-quality A. incana litter and that the macro-detritivores tested differed in their decomposition-stimulating effects, with earthworms having the largest influence. Decomposition processes increased with increasing number of macro-detritivore species, and positive net diveristy effects occurred in several macro-detritivore treatments. However, after correction for macro-detritivore biomass, all interspecific differences in macro-detritivore effects, as well as the positive effects of species number on subarctic litter decomposition disappeared. The net diversity effects also appeared to be driven by variation in biomass, with a possible exception of net diversity effects in mass loss. Based on these results, we conclude that the expected climate change-induced range expansion of macro-detritivores into subarctic regions is likely to result in accelerated decomposition rates. Our results also indicate that the magnitude of macro-detritivore effects on subarctic decomposition will mainly depend on macro-detritivore biomass, rather than on macro-detritivore species number or identity. [ABSTRACT FROM AUTHOR]

The mass loss of litter mixtures is often different than expected based on the mass loss of the component species. We investigated if the identity of neighbour species affects these litter-mixing effects. To achieve this, we compared decomposition rates in monoculture and in all possible two-species combinations of eight tree species, widely differing in litter chemistry, set out in two contrasting New Zealand forest types. Litter from the mixed-species litter bags was separated into its component species, which allowed us to quantify the importance of litter-mixing effects and neighbour identity, relative to the effects of species identity, litter chemistry and litter incubation environment. Controlling factors on litter decomposition rate decreased in importance in the order: species identity (litter quality) >> forest type >> neighbour species. Species identity had the strongest influence on decomposition rate. Interspecific differences in initial litter lignin concentration explained a large proportion of the interspecific differences in litter decomposition rate. Litter mass loss was higher and litter-mixture effects were stronger on the younger, more fertile alluvial soils than on the older, less-fertile marine terrace soils. Litter-mixture effects only shifted percentage mass loss within the range of 1.5%. There was no evidence that certain litter mixtures consistently showed interactive effects. Contrary to common theory, adding a relatively fast-decomposing species generally slowed down the decomposition of the slower decomposing species in the mixture. This study shows that: (1) species identity, litter chemistry and forest type are quantitatively the most important drivers of litter decomposition in a New Zealand rain forest; (2) litter-mixture effects—although statistically significant—are far less important and hardly depend on the identity and the chemical characteristics of the neighbour species; (3) additive effects predominate in this ecosystem, so that mass dynamics of the mixtures can be predicted from the monocultures. [ABSTRACT FROM AUTHOR]

Worldwide decomposition rates depend both on climate and the legacy of plant functional traits as litter quality. To quantify the degree to which functional differentiation among species affects their litter decomposition rates, we brought together leaf trait and litter mass loss data for 818 species from 66 decomposition experiments on six continents. We show that: (i) the magnitude of species-driven differences is much larger than previously thought and greater than climate-driven variation; (ii) the decomposability of a species’ litter is consistently correlated with that species’ ecological strategy within different ecosystems globally, representing a new connection between whole plant carbon strategy and biogeochemical cycling. This connection between plant strategies and decomposability is crucial for both understanding vegetation–soil feedbacks, and for improving forecasts of the global carbon cycle. [ABSTRACT FROM AUTHOR]

Although hemiparasitic plants have a number of roles in shaping the structure and composition of plant communities, the impact of this group on ecosystem processes, such as decomposition and nutrient cycling, has been poorly studied. In order to better understand the potential role of hemiparasites in these processes, a comparison of leaf and litter tissue quality, nitrogen (N) resorption, and decomposability with those of a wide range of other plant groups (involving a total of 72 species and including other groups with access to alternative nutrient sources, such as nitrogen fixers and carnivorous plants) was undertaken in several sub-arctic habitats. The foliar N concentration of hemiparasites generally exceeded that of co-occurring species. Further, hemiparasites (and N fixers) exhibited lower N resorption efficiencies than their counterparts with no major alternative N source. As a consequence, annual and perennial hemiparasite litter contained, on average, 3.1% and 1.9% N, respectively, compared with 0.77-1.1% for groups without a major alternative N source. Hemiparasite litter lost significantly more mass during decomposition than many, but not all, co-occurring species. These results were combined with those of a litter trapping experiment to assess the potential impact of hemiparasites on nutrient cycling. The common sub-arctic hemiparasite Bartsia alpina was estimated to increase the total annual N input from litter to the soil by ∼42% within 5 cm of its stems, and by ∼53% across a site with a Bartsia alpina stem density of 43 stems/m[sup2]. Our results therefore provide clear evidence in favor of a novel mechanism by which hemiparasites (in parallel with N-fixing species) may influence ecosystems in which they occur. [ABSTRACT FROM AUTHOR]