Litter decomposition of a pine plantation is affected by species evenness and soil nitrogen availability

Background and aims

Litter decomposition is a key process controlling flows of energy and nutrients in ecosystems. Altered biodiversity and nutrient availability may affect litter decomposition. However, little is known about the response of litter decomposition to co-occurring changes in species evenness and soil nutrient availability.

Methods

We used a microcosm experiment to evaluate the simultaneous effects of species evenness (two levels), identity of the dominant species (three species) and soil N availability (control and N addition) on litter decomposition in a Mongolian pine (Pinus sylvestris var. mongolica) plantation in Northeast China. Mongolian pine needles and senesced aboveground materials of two dominant understory species (Setaria viridis and Artemisia scoparia) were used for incubation.

Results

Litter evenness, dominant species identity and N addition significantly affected species interaction and litter decomposition. Higher level of species evenness increased the decomposition rate of litter mixtures and decreased the incidence of antagonistic effects. A. scoparia-dominated litter mixtures decomposed faster than P. sylvestris var. mongolica- and S. viridis-dominated litter mixtures. Notably, N addition increased decomposition rate of both single-species litters and litter mixtures, and meanwhile altered the incidence and direction of non-additive effects during decomposition of litter mixtures. The presence of understory species litters stimulated the decomposition rate of pine litters irrespective of N addition, whereas the presence of pine litters suppressed the mass loss of A. scoparia litters. Moreover, N addition weakened the promoting effects of understory species litters on decomposition of pine litters.

Conclusions

Pine litter retarded the decomposition of understory species litters whereas its own decomposition was accelerated in mixtures. Nitrogen addition and understory species evenness altered species interaction through species-specific responses in litter mixtures and thus affected litter decomposition in Mongolian pine forests, which could produce a potential influence on ecosystem C budget and nutrient cycling.     

Mixing Effects of Understory Plant Litter on Decomposition and Nutrient Release of Tree Litter in Two Plantations in Northeast China

Understory vegetation plays a crucial role in carbon and nutrient cycling in forest ecosystems; however, it is not clear how understory species affect tree litter decomposition and nutrient dynamics. In this study, we examined the impacts of understory litter on the decomposition and nutrient release of tree litter both in a pine (Pinus sylvestris var. mongolica) and a poplar (Populus × xiaozhuanica) plantation in Northeast China. Leaf litter of tree species, and senesced aboveground materials from two dominant understory species, Artemisia scoparia and Setaria viridis in the pine stand and Elymus villifer and A. sieversiana in the poplar stand, were collected. Mass loss and N and P fluxes of single-species litter and three-species mixtures in each of the two forests were quantified. Data from single-species litterbags were used to generate predicted mass loss and N and P fluxes for the mixed-species litterbags. In the mixture from the pine stand, the observed mass loss and N release did not differ from the predicted value, whereas the observed P release was greater than the predicted value. However, the presence of understory litter decelerated the mass loss and did not affect N and P releases from the pine litter. In the poplar stand, litter mixture presented a positive non-additive effect on litter mass loss and P release, but an addition effect on N release. The presence of understory species accelerated only N release of poplar litter. Moreover, the responses of mass loss and N and P releases of understory litter in the mixtures varied with species in both pine and poplar plantations. Our results suggest that the effects of understory species on tree litter decomposition vary with tree species, and also highlight the importance of understory species in litter decomposition and nutrient cycles in forest ecosystems.     

Do non‐additive effects on decomposition in litter‐mix experiments result from differences in resource quality between litters?

Differences in resource quality between litter species have been postulated to explain why litter-mixtures may decompose at a different rate to that which would be predicted from single species litters (termed 'non-additive effects'). In particular, positive, non-additive effects of litter-mixing on decomposition have been explained by differences in initial nitrogen concentration between litter species. This interpretation is confounded because litter species that differ in nitrogen content also differ by a number of other resource quality attributes. Thus, to investigate whether initial nitrogen concentration does account for positive, non-additive effects of litter-mixing, we mixed grass litters that differed in initial nitrogen concentration but not species or structural plant part identity, and then followed mass loss from the litter-mixes over time. We used the litterbag technique and three grass species for which a gradient of four distinct initial nitrogen concentrations had been generated. We produced all no- to four-mix compositions of litter qualities for each species. Litter from different species was never mixed.
Contrary to what would be predicted, we found that when litters of the same species but with different initial nitrogen concentrations were mixed, that negative, non-additive effects on decomposition were generally observed. In addition, we found that once mixed, increasing litter quality richness from two to four mixtures had no significant, non-additive effect on decomposition. Litter quality composition explained little of the experimental variation when compared to litter quality richness, and different compositions generally behaved in the same manner. Our findings challenge the commonly held assumption that differences in nitrogen concentration between plant species are responsible for positive, non-additive effects of litter-mixing on decomposition.     

Decomposition and nutrient dynamics in mixed litter of Mediterranean species

In the last decade a great research effort addressed the effects of litter diversity on ecosystem functions, reporting both synergistic and antagonistic effects for decomposition dynamics. Four coexisting Mediterranean species, representing a range of litter quality, were used to arrange litter mixtures at three diversity levels for a litterbag decomposition experiment. Species identity appeared as the major determinant for litter mass loss (Coronilla emerus～Hedera helix>Festuca drymeia>Quercus ilex) and nutrient release, with rates for all leaf litter types following the sequence K>N>Mg≥Ca>>Fe. Additive diversity effects were prevalent pooling together all data but also for nutrients separately. Antagonistic interactions were more common than synergistic in the cases of mass loss, N and Ca contents, but not for K, Mg and Fe dynamics. The number of species in the litterbag significantly affected the outcome of non-additive interactions, which were mostly antagonistic for two-species mixtures, and synergistic for the combined 4 species. Litter quality appears to be the most important factor affecting mass loss and nutrient dynamics, while litter diversity, influencing the rates of these processes, plays an important role in reducing their variability, thus suggesting a greater stability of ecosystems properties in presence of mixed litter.     

Non-additive effects of litter mixtures on decomposition of leaf litters in a Mediterranean maquis

Many studies across a range of ecosystems have shown that decomposition in mixed litter is not predictable from single-species results due to synergistic or antagonistic interactions. Some studies also reveal that species composition and relative abundance may be more important than just richness in driving non-additive effects. Most studies on litter decomposition in Mediterranean maquis, an high-diversity shrubby ecosystem, have dealt exclusively with single species. In this study we investigated, at the individual-litter level, as well as at the litter-mixture level, the effect of litter mixing on decomposition of 3-species litter assemblages with different relative abundance of the component litters; we set up two types of litter assemblages that reflected the heterogeneity of bush cover in the inner maquis and at the edge maquis/gaps, as related to the leaf traits, i.e. sclerophylly vs mesophylly. We measured mass loss, decay of lignin, cellulose and ADSS (acid detergent soluble substances) and fungal mycelium ingrowth. The results show that over a 403-day incubation period, the decomposition of individual litters in mixtures deviated from that of monospecific litters and had different directions. In litter mixtures of the sclerophylls Phillyrea angustifolia and Pistacea lentiscus with the mesophyll Cistus, decomposition was lower than expected (antagonistic effect); in the mixtures of litters with similar physical structure (Ph. angustifolia and P. lentiscus with Quercus ilex) decomposition was faster than expected (synergistic effect). When considering the different decomposition phases, both negative and positive effects occurred in Quercus mixtures depending on the phase of decomposition. In both types of 3-species litter assemblages the greatest effect occurred in uneven mixtures rather than in even mixtures. Our results show that species composition drives the direction whilst the decomposability and the relative abundance drive the magnitude of non-additive effects of litter mixing on decomposition.     

Non‐additive effects of invasive tree litter shift seasonal N release: a potential invasion feedback

Many invasive plant species strongly alter ecosystem processes by producing leaf litter that decomposes faster and releases N more quickly than that of native species. However, while most studies of invasive species litter impacts have only considered the decomposition of species in monoculture, forest litter layers typically contain litter from many species. Many litter mixtures decompose in a non‐additive manner, in which the mixture decomposes more quickly (synergistic effect) or more slowly (antagonistic effect) than would be expected based on decomposition of the component species’ litters in isolation. We investigated the potential for non‐additive effects of invasive species’ litter by conducting a one‐year litter bag experiment in which we mixed the litters of four native tree species with each of four invasive species. Litter mixtures frequently lost mass at non‐additive rates, although not at every loading ratio, and the presence, sign, and strength of effects depended on species composition. Non‐additive effects on N loss occurred in more litter combinations, and were almost always antagonistic at 90 days and synergistic at 365 days. Invasive species litter with lower C:N led to more strongly synergistic N loss with time. During the growing season, non‐additive patterns of N loss almost always resulted in increased N release – up to six times greater than would be expected based on single‐species decomposition. Consequently, we suggest that invasive species may further synchronize N release from the litter layer with plant N demand, enhancing any positive litter feedback to invasion. These results highlight the need to consider non‐additive effects of litter mixing in invaded forest communities, and suggest that estimates of invasive species’ impacts on ecosystem processes would be improved by considering these effects.     

Tree species richness affects litter production and decomposition rates in a tropical biodiversity experiment

We report data on leaf litter production and decomposition from a manipulative biodiversity experiment with trees in tropical Panama, which has been designed to explore the relationship between tree diversity and ecosystem functioning. A total of 24 plots (2025 m²) were established in 2001 using six native tree species, with 1‐, 3‐, and 6‐species mixtures. We estimated litter production during the dry season 2005 with litter traps; decomposition was assessed with a litter bag approach during the following wet season. Litter production during the course of the dry season was highly variable among the tree species. Tree diversity significantly affected litter production, and the majority of the intermediate diverse mixtures had higher litter yields than expected based on yields in monoculture. In contrast, high diverse mixtures did not show such overyielding in litter production. Litter decomposition rates were also highly species‐specific, and were related to various measures of litter quality (C/N, lignin/N, fibre content). We found no overall effect of litter diversity if the entire litter mixtures were analyzed, i.e. mixing species resulted in pure additive effects and observed decomposition rates were not different from expected rates. However, the individual species changed their decomposition pattern depending on the diversity of the litter mixture, i.e. there were species‐specific responses to mixing litter. The analysis of temporal C and N dynamics within litter mixtures gave only limited evidence for nutrient transfer among litters of different quality. At this early stage of our tree diversity experiment, there are no coherent and general effects of tree species richness on both litter production and decomposition. Within the scope of the biodiversity‐ecosystem functioning relationship, our results therefore highlight the process‐specific effects diversity may have. Additionally, species‐specific effects on ecosystem processes and their temporal dynamics are important, but such effects may change along the gradient of tree diversity.     

Mass loss and nutrient dynamics during litter decomposition under three mixing treatments in a typical steppe in Inner Mongolia

Background and aims

Mixing effects during litter decomposition could occur between two or more different litter species because of the potential nutrient transfer. However, evidence of mixing effects is variable and the underlying mechanisms remain unclear. Using a three-year decomposition experiment, we aim to examine for the effects of litter mixing and position on decomposition rates and nitrogen (N) and phosphorus (P) dynamics.

Methods

We studied litter decomposition of Stipa krylovii (Sk) and Astragalus galactites (Ag), two dominant species with contrasting litter quality, in a typical steppe of northern China in both single decomposition and three mixing treatments. The three mixing treatments included thorough mixing (Sk-Ag), Ag over Sk (Ag/Sk), and Sk over Ag (Sk/Ag).

Results

Both the Sk-Ag and the Sk/Ag mixture had negative mixing effects on the mass loss of the litter mixture, while the Ag/Sk mixture had a neutral mixing effect. The percent mass loss was higher when the litter species was placed at the top (25.0 and 51.9 % of mass remaining for Ag and Sk, respectively) than at the bottom (38.3 and 61.8 % of mass remaining for Ag and Sk, respectively). The Sk/Ag mixture had negative effects on the release of N while all three mixing treatments had positive effects on the release of P.

Conclusions

Our results indicate that: (1) mixing treatments can induce different mixing effects; (2) environmental factors likely play an important role in controlling the mixing effect; and (3) litter-mixtures have different non-additive effects on N and P, which may further increase the heterogeneity of N and P availability as the two litter species may fall differentially in terms of space and time.     

Nitrogen and Phosphorus Release from Mixed Litter Layers is Lower than Predicted from Single Species Decay

Ecosystem-level nutrient dynamics during decomposition are often estimated from litter monocultures. If species effects are additive, we can statistically predict nutrient dynamics in multi-species systems from monoculture work, and potential consequences of species loss. However, if species effects are dependent on interactions with other litter species (that is, non-additive), predictions based on monoculture data will likely be inaccurate. We conducted a 3-year, full-factorial, mixed-litter decomposition study of four dominant tree species in a temperate forest and measured nitrogen and phosphorus dynamics to explore whether nutrient dynamics in mixtures were additive or non-additive. Following common approaches, we used litterfall data to predict nutrient dynamics at the ecosystem-level. In mixtures, we observed non-additive effects of litter mixing on nutrient dynamics: the presence of nutrient-rich species in mixture facilitated nutrient release, whereas nutrient-poor species facilitated nutrient retention. Fewer nutrients were released from mixtures containing high-quality litter, and more immobilized from mixtures containing low-quality litter, than predicted from monocultures, creating a difference in overall nutrient release between predicted and actual dynamics in litter mixtures. Nutrient release at the ecosystem-level was greatly overestimated when based on monocultures because the effect of species interactions on nutrient immobilization was not accounted for. Our data illustrate that the identity of species in mixtures is key to their role in non-additive interactions, with repercussions for mineral nutrient availability and storage. These results suggest that predictions of ecosystem-level nutrient dynamics using litter monoculture data likely do not accurately represent actual dynamics because the effects of litter species interactions are not incorporated. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Long-term presence of tree species but not chemical diversity affect litter mixture effects on decomposition in a neotropical rainforest

Plant litter diversity effects on decomposition rates are frequently reported, but with a strong bias towards temperate ecosystems. Altered decomposition and nutrient recycling with changing litter diversity may be particularly important in tree species-rich tropical rainforests on nutrient-poor soils. Using 28 different mixtures of leaf litter from 16 Amazonian rainforest tree species, we tested the hypothesis that litter mixture effects on decomposition increase with increasing functional litter diversity. Litter mixtures and all single litter species were exposed in the field for 9 months using custom-made microcosms with soil fauna access. In order to test the hypothesis that the long-term presence of tree species contributing to the litter mixtures increases mixture effects on decomposition, microcosms were installed in a plantation at sites including the respective tree species composition and in a nearby natural forest where these tree species are absent. We found that mixture decomposition deviated from predictions based on single species, with predominantly synergistic effects. Functional litter diversity, defined as either richness, evenness, or divergence based on a wide range of chemical traits, did not explain the observed litter mixture effects. However, synergistic effects in litter mixtures increased with the long-term presence of tree species contributing to these mixtures as the home field advantage hypothesis assumes. Our data suggest that complementarity effects on mixed litter decomposition may emerge through long-term interactions between aboveground and belowground biota.     

C,N and P fertilization in an Amazonian rainforest supports stoichiometric dissimilarity as a driver of litter diversity effects on decomposition

Plant leaf litter generally decomposes faster as a group of different species than when individual species decompose alone, but underlying mechanisms of these diversity effects remain poorly understood. Because resource C : N : P stoichiometry (i.e. the ratios of these key elements) exhibits strong control on consumers, we supposed that stoichiometric dissimilarity of litter mixtures (i.e. the divergence in C : N : P ratios among species) improves resource complementarity to decomposers leading to faster mixture decomposition. We tested this hypothesis with: (i) a wide range of leaf litter mixtures of neotropical tree species varying in C : N : P dissimilarity, and (ii) a nutrient addition experiment (C, N and P) to create stoichiometric similarity. Litter mixtures decomposed in the field using two different types of litterbags allowing or preventing access to soil fauna. Litter mixture mass loss was higher than expected from species decomposing singly, especially in presence of soil fauna. With fauna, synergistic litter mixture effects increased with increasing stoichiometric dissimilarity of litter mixtures and this positive relationship disappeared with fertilizer addition. Our results indicate that litter stoichiometric dissimilarity drives mixture effects via the nutritional requirements of soil fauna. Incorporating ecological stoichiometry in biodiversity research allows refinement of the underlying mechanisms of how changing biodiversity affects ecosystem functioning.     

Limited evidence that mixing leaf litter accelerates decomposition or increases diversity of decomposers in streams of eastern Canada

Most studies of terrestrial litter decomposition in streams and rivers have used leaves from a single tree species, but leaf packs in streams in eastern North America are usually mixtures of two or more species. Litter mixtures may decay more quickly than either of the component species. If so, estimates of stream energy and nutrient budgets may be inaccurate. In northern Nova Scotia, Canada, we measured mass loss from binary mixtures (1:1 mass ratio) of leaf litter in mesh bags, using freshly fallen or air-dried litter from five species of canopy trees. We repeated the experiment eight times, in summer and fall, in two streams and a small river, over 3 years. In some trials we enumerated benthic invertebrate and fungal colonization of decaying litter. Although there were marked differences in mass loss rates among litter types, decomposition was accelerated in mixtures relative to the mean of the component species in only three of eight trials, and only in mixtures containing N-rich speckled alder leaves. Mixing yellow birch and red maple leaves inhibited decomposition. Diversity (Shannon–Weaver Index), species richness, and abundance of aquatic hyphomycete fungi, as indexed by conidial production, were never greater (and sometimes less) on litter mixtures than on the component species. Total numbers, taxonomic richness and diversity of benthic invertebrates generally, and litter-feeding species in particular, were not augmented by mixing litter types. Litter mixtures appear to dilute a preferred substrate with patches of a less preferred substrate. Our results provide only weak support for the contention that combining two litter types leads to acceleration of decomposition rates. Handling editor: K. Martens     

凋落物多样性及组成对凋落物分解和土壤微生物群落的影响

凋落物分解是生态系统营养物质循环的核心过程,而土壤微生物群落在凋落物分解过程中扮演着极其重要且不可替代的角色。随着生物多样性的丧失日益严峻,探讨凋落物多样性及组成对凋落物分解和土壤微生物群落的影响,不仅有助于了解凋落物分解的内在机制,而且可为退化草原生态系统的恢复提供参考。以内蒙古呼伦贝尔草原退化恢复群落中的草本植物为研究对象,依据植物多度、盖度、频度和物种的重要值及其在群落中的恢复程度筛选出排序前4的羊草（Leymus chinensis）、茵陈蒿（Artemisia capillaris）、麻花头（Serratula centauroides）、二裂委陵菜（Potentilla bifurca）的凋落物为实验材料,通过设置3种凋落物多样性水平（1,2,4）,包括11种凋落物组合（单物种凋落物共4种,两物种凋落物混合共6种,四物种凋落物混合共1种）,利用磷脂脂肪酸（PLFA）方法来研究分解60 d后凋落物多样性及组成对凋落物分解和土壤微生物群落的影响。结果表明：（1）凋落物物种多样性仅对C残余率具有显著影响,表现在两物种混合凋落物C残余率显著低于单物种凋落物,而凋落物组成对所观测的4个凋落物分解参数（质量、C、N残余率以及C/N）均具有显著影响;（2）凋落物物种多样性对细菌（B）含量具有显著影响,而凋落物组成对真菌（F）含量具有显著影响,两者对F/B以及微生物总量均无显著影响;（3）冗余分析结果表明凋落物组成与凋落物分解相关指标（凋落物质量、C、N残余率及C/N）和土壤微生物（真菌、细菌含量）的相关关系高于凋落物多样性。（4）进一步建立结构方程模型（Structural Equation Model,SEM）发现,凋落物初始C含量对凋落物质量、C、N残余率及C/N有显著正的直接影响;凋落物木质素含量对凋落物质量、C、N残余率有显著正的直接影响;凋落物初始N含量对N残余率有显著正的直接影响,而对C残余率及C/N有显著负的直接影响;凋落物初始C/N对凋落物质量、N残余率有显著正的直接影响,而对C/N有显著负的直接影响。此外,凋落物初始C、N、木质素含量及C/N均对真菌含量具有显著正影响,并可通过真菌对凋落物质量分解产生显著负的间接影响。以上结果表明该退化恢复区域优势种凋落物分解以初始C、木质素为主导,主要通过土壤真菌影响凋落物的分解进程,这将减缓凋落物的分解速率进而减慢草原生态系统的进程。这些结果为凋落物多样性及组成对自身分解和土壤微生物群落的影响提供了实验依据,也为进一步分析凋落物分解内在机制以及草原生态系统的恢复提供了数据参考。     

Neighbour identity hardly affects litter-mixture effects on decomposition rates of New Zealand forest species

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.     

Does mixing litter of different qualities alter stream microbial diversity and functioning on individual litter species?

We examined effects of leaf litter quality and species mixing on microbial community diversity and litter processing in a forested headwater stream. Single- and mixed-species litter from dominant tree species ( Liriodendron tulipifera , Acer rubrum , Quercus prinus , Rhododendron maximum ) were incubated in a southern Appalachian headwater stream. Litter carbon-to-nitrogen ratios (C:N), mass loss, microbial respiration, and microbial community diversity were analyzed on individual litter species after incubation. Initial C:N varied widely among individual litter species, and these differences persisted throughout the 50-day incubation period. Litter C:N of the recalcitrant species R. maximum remained higher than that of all other litter species, and C:N of R. maximum and L. tulipifera increased when both species were present together in a mixture. Although mass loss of individual species was generally unaffected by mixing, microbial respiration was greater on A. rubrum and Q. prinus litter incubated with R. maximum compared to either species alone. Enhanced resource heterogeneity, which was experimentally achieved by litter mixing low- and higher-quality litter species, resulted in apparent shifts in microbial community diversity on individual litter species. Responses of bacterial and fungal community diversity to litter mixing varied among individual litter species. Our results suggest that changes in tree species composition in riparian forests and subsequent changes in litter resource heterogeneity could alter stream microbial community diversity and function. As bacteria and fungi are important decomposers of plant litter in aquatic ecosystems, resource-dependent changes in microbial communities could alter detrital processing dynamics in streams.     

Nitrogen addition stimulates forest litter decomposition and disrupts species interactions in Patagonia,Argentina

Nitrogen (N) deposition and biodiversity loss are important drivers of global change, with uncertain consequences for carbon (C) and nutrient turnover in terrestrial ecosystems. We evaluated the simultaneous effects of N deposition and plant diversity on litter decomposition within a temperate forest in Patagonia. We identified ‘tree triangles’ created by the intersection of three tree‐canopies that directly controlled micro‐environmental conditions on the forest floor, and combined it with an N addition treatment. Triangles were composed of one or three Nothofagus species (N. dombeyi, N. obliqua or N. nervosa). We placed litterbags containing litter of each of the Nothofagus species and litterbags containing a mixture of the three species within all triangles and assessed mass loss over 2 years. We used a standard litter type in all triangles to independently evaluate triangle effects on decomposition. N addition had strong and positive effects on decomposition with an average 46% increase in the decomposition constant. Litter species significantly differed in their response to N addition; litter with higher lignin content and lower labile C content had larger increase in decomposition due to fertilization. Also, N addition disrupted two types of species interactions that control decomposition. The affinity relation between litter and decomposers, that enhanced decomposition of home litter (‘home‐field advantage’) that was demonstrated to be significant for all three Nothofagus species, disappeared with N addition. Second, N addition modified litter species interactions, transforming neutral effects of litter mixtures to positive, nonadditive effects on mass loss. Finally, N addition stimulated N release from decomposing litter which was modulated by plant species effects. Together, these results suggest that N addition to unpolluted forests increases C loss, contrary to what has been observed for temperate forests in industrialized areas of the world, and that alterations in nutrient pools have effects on species interactions, including the elimination of affinity effects for decomposition.     

Litter evenness influences short-term peatland decomposition processes

There is concern that changes in climate and land use could increase rates of decomposition in peatlands, leading to release of stored C to the atmosphere. Rates of decomposition are driven by abiotic factors such as temperature and moisture, but also by biotic factors such as changes in litter quality resulting from vegetation change. While effects of litter species identity and diversity on decomposition processes are well studied, the impact of changes in relative abundance (evenness) of species has received less attention. In this study we investigated effects of changes in short-term peatland plant species evenness on decomposition in mixed litter assemblages, measured as litter weight loss, respired CO₂ and leachate C and N. We found that over the 307-day incubation period, higher levels of species evenness increased rates of decomposition in mixed litters, measured as weight loss and leachate dissolved organic N. We also found that the identity of the dominant species influenced rates of decomposition, measured as weight loss, CO₂ flux and leachate N. Greatest rates of decomposition were when the dwarf shrub Calluna vulgaris dominated litter mixtures, and lowest rates when the bryophyte Pleurozium schreberi dominated. Interactions between evenness and dominant species identity were also detected for litter weight loss and leachate N. In addition, positive non-additive effects of mixing litter were observed for litter weight loss. Our findings highlight the importance of changes in the evenness of plant community composition for short-term decomposition processes in UK peatlands.     

Leaf Litter Mixtures Alter Microbial Community Development: Mechanisms for Non-Additive Effects in Litter Decomposition

To what extent microbial community composition can explain variability in ecosystem processes remains an open question in ecology. Microbial decomposer communities can change during litter decomposition due to biotic interactions and shifting substrate availability. Though relative abundance of decomposers may change due to mixing leaf litter, linking these shifts to the non-additive patterns often recorded in mixed species litter decomposition rates has been elusive, and links community composition to ecosystem function. We extracted phospholipid fatty acids (PLFAs) from single species and mixed species leaf litterbags after 10 and 27 months of decomposition in a mixed conifer forest. Total PLFA concentrations were 70% higher on litter mixtures than single litter types after 10 months, but were only 20% higher after 27 months. Similarly, fungal-to-bacterial ratios differed between mixed and single litter types after 10 months of decomposition, but equalized over time. Microbial community composition, as indicated by principal components analyses, differed due to both litter mixing and stage of litter decomposition. PLFA biomarkers a15∶0 and cy17∶0, which indicate gram-positive and gram-negative bacteria respectively, in particular drove these shifts. Total PLFA correlated significantly with single litter mass loss early in decomposition but not at later stages. We conclude that litter mixing alters microbial community development, which can contribute to synergisms in litter decomposition. These findings advance our understanding of how changing forest biodiversity can alter microbial communities and the ecosystem processes they mediate.     

Greater accumulation of litter in spruce (Picea abies) compared to beech (Fagus sylvatica) stands is not a consequence of the inherent recalcitrance of needles

Background and aims

Replacement of beech by spruce is associated with changes in soil acidity, soil structure and humus form, which are commonly ascribed to the recalcitrance of spruce needles. It is of practical relevance to know how much beech must be admixed to pure spruce stands in order to increase litter decomposition and associated nutrient cycling. We addressed the impact of tree species mixture within forest stands and within litter on mass loss and nutritional release from litter.

Methods

Litter decomposition was measured in three adjacent stands of pure spruce (Picea abies), mixed beech-spruce and pure beech (Fagus sylvatica) on three nutrient-rich sites and three nutrient-poor sites over a three-year period using the litterbag method (single species and mixed species bags).

Results

Mass loss of beech litter was not higher than mass loss of spruce litter. Mass loss and nutrient release were not affected by litter mixing. Litter decay indicated non-additive patterns, since similar remaining masses under pure beech (47%) and mixed beech-spruce (48%) were significantly lower than under pure spruce stands (67%). Release of the main components of the organic substance (C_org, N_tot, P, S, lignin) and associated K were related to mass loss, while release of other nutrients was not related to mass loss.

Conclusions

In contradiction to the widely held assumption of slow decomposition of spruce needles, we conclude that accumulation of litter in spruce stands is not caused by recalcitrance of spruce needles to decay; rather adverse environmental conditions in spruce stands retard decomposition. Mixed beech-spruce stands appear to be as effective as pure beech stands in counteracting these adverse conditions.     

Non-Additive Effects on Decomposition from Mixing Litter of the Invasive Mikania micrantha H.B.K. with Native Plants

A common hypothesis to explain the effect of litter mixing is based on the difference in litter N content between mixed species. Although many studies have shown that litter of invasive non-native plants typically has higher N content than that of native plants in the communities they invade, there has been surprisingly little study of mixing effects during plant invasions. We address this question in south China where Mikania micrantha H.B.K., a non-native vine, with high litter N content, has invaded many forested ecosystems. We were specifically interested in whether this invader accelerated decomposition and how the strength of the litter mixing effect changes with the degree of invasion and over time during litter decomposition. Using litterbags, we evaluated the effect of mixing litter of M. micrantha with the litter of 7 native resident plants, at 3 ratios: M₁ (1∶4, = exotic:native litter), M₂ (1∶1) and M₃ (4∶1, = exotic:native litter) over three incubation periods. We compared mixed litter with unmixed litter of the native species to identify if a non-additive effect of mixing litter existed. We found that there were positive significant non-additive effects of litter mixing on both mass loss and nutrient release. These effects changed with native species identity, mixture ratio and decay times. Overall the greatest accelerations of mixture decay and N release tended to be in the highest degree of invasion (mix ratio M₃) and during the middle and final measured stages of decomposition. Contrary to expectations, the initial difference in litter N did not explain species differences in the effect of mixing but overall it appears that invasion by M. micrantha is accelerating the decomposition of native species litter. This effect on a fundamental ecosystem process could contribute to higher rates of nutrient turnover in invaded ecosystems.