Tree litter functional diversity and nitrogen concentration enhance litter decomposition via changes in earthworm communities

Biodiversity is a major driver of numerous ecosystem functions. However, consequences of changes in forest biodiversity remain difficult to predict because of limited knowledge about how tree diversity influences ecosystem functions. Litter decomposition is a key process affecting nutrient cycling, productivity, and carbon storage and can be influenced by plant biodiversity. Leaf litter species composition, environmental conditions, and the detritivore community are main components of the decomposition process, but their complex interactions are poorly understood. In this study, we tested the effect of tree functional diversity (FD) on litter decomposition in a field experiment manipulating tree diversity and partitioned the effects of litter physiochemical diversity and the detritivore community. We used litterbags with different mesh sizes to separate the effects of microorganisms and microfauna, mesofauna, and macrofauna and monitored soil fauna using pitfall traps and earthworm extractions. We hypothesized that higher tree litter FD accelerates litter decomposition due to the availability of complementary food components and higher activity of detritivores. Although we did not find direct effects of tree FD on litter decomposition, we identified key litter traits and macrodetritivores that explained part of the process. Litter mass loss was found to decrease with an increase in leaf litter carbon:nitrogen ratio. Moreover, litter mass loss increased with an increasing density of epigeic earthworms, with most pronounced effects in litterbags with a smaller mesh size, indicating indirect effects. Higher litter FD and litter nutrient content were found to increase the density of surface‐dwelling macrofauna and epigeic earthworm biomass. Based on structural equation modeling, we conclude that tree FD has a weak positive effect on soil surface litter decomposition by increasing the density of epigeic earthworms and that litter nitrogen‐related traits play a central role in tree composition effects on soil fauna and decomposition.     

氮沉降对森林凋落物分解的影响

氮沉降增加作为全球变化的重要现象之一,已经并将继续对森林凋落物分解产生影响.综述了国内外氮沉降对森林凋落物分解影响及其机理的研究现状.氮沉降对凋落物分解的影响可分为直接影响和间接影响.氮沉降通过影响森林地被物组成和凋落物化学成分,间接影响凋落物分解.氮沉降对凋落物分解的直接影响表现为促进、无影响和抑制3种效果.分析了产生以上影响效果的作用机理,介绍了氮沉降对森林凋落物分解影响的研究方法,探讨了目前研究存在的问题,讨论了未来该方面研究的重点和方向.     

桂西北喀斯特区原生林与次生林凋落叶降解和养分释放

凋落叶降解及养分释放研究对喀斯特生态脆弱区森林生态系统的恢复与重建具有重要指导意义。本文选取桂西北喀斯特区3种原生林与3种次生林进行比较,研究其凋落叶降解与降解过程中的营养元素释放规律以及降解速率的影响因子。结果表明,原生林凋落叶的降解速率略大于次生林。C、N、K元素在前180天释放速率较快,随后趋于稳定。次生林凋落叶总P含量在降解初始阶段呈净积累,随后净释放,而原生林的凋落叶在降解360天后仍呈现P素净积累。相关分析表明,凋落叶降解速率与凋落叶初始总N、木质素含量及木质素:N比值呈负相关,与C:N比呈正相关。综合比较发现,次生林圆叶乌桕(Sapium rotundifolium Hemsl)凋落叶的降解速率与养分释放速率较快,是喀斯特退化土地及植被恢复过程中潜在的优势种和建群种。     

Subtropical montane tree litter decomposition: Links with secondary forest types and species' shade tolerance

Abstract Litter decomposition plays an important role in secondary forest recovery in the tropics. In this study we assessed the decomposition rates of tree litter in species from different secondary forest types and with different shade tolerances. The three secondary forest types analysed are related to the effects of different previous land use intensities. The typical forest type (TYP) is related to low land use intensity, Alnus acuminata‐dominated forest (ALN) type is related to medium land‐use intensity and Amomyrtella güili‐dominated forest type (AMO) is related to high land use intensity. The effect of shade tolerance was assessed using maximum height of each species as an indicator of its light requirements. Associations with leaf functional traits such as specific leaf area (SLA), and tensile strength (LTS) were also assessed. We found that leaves of species from the TYP forest type decompose faster than those of the ALN and AMO forest types. These changes were consistent with differences in the SLA of the species, which was higher in the TYP forest type than in the ALN and AMO forest types. SLA, LTS and decomposition were not significantly correlated with tree maximum height. Our results show that the secondary forest types, which are related to land use intensities prior to abandonment have an important influence on litter decomposition. This implies potential long‐term effects on soil properties and species composition.     

Inter‐specific Variation in Foliar Nutrients and Resorption of Nine Canopy‐tree Species in a Secondary Neotropical Rain Forest

The influence of environmental gradients on the foliar nutrient economy of forests has been well documented; however, we have little understanding of what drives variability among individuals within a single forest stand, especially tropical forests. We evaluated inter‐ and intra‐specific variation in nutrient resorption, foliar nutrient concentrations and physical leaf traits of nine canopy tree species within a 1‐ha secondary tropical rain forest in northeastern Costa Rica. Both nitrogen (N) and phosphorus (P) resorption efficiency (RE) and proficiency of the nine tree species varied significantly among species, but not within. Both N and P RE were significantly negatively related to leaf specific strength. Green leaf N and P concentrations were strongly negatively related to leaf mass per area, and senesced leaf nutrient concentrations were significantly positively related to green leaf nutrient concentrations. This study reveals a strong influence of physical leaf traits on foliar nutrient and resorption traits of co‐occurring species in a secondary wet tropical forest stand.     

Plant species control and soil faunal involvement in the processes of above‐ and below‐ground litter decomposition

Among the factors determining litter decomposition rates, the role of soil fauna as decomposers still remains unclear, especially for how they are involved in decomposing below‐ground root litter compared to their relatively‐known contributions to decomposing above‐ground leaf litter. We conducted a litterbag experiment using two sizes of meshes and pursued the leaf and root decomposition of six major tree species in a Japanese temperate forest over 411‐days to test the interactive effects of soil mesofauna and litter quality addressed based on two features (litter types and species) on the process. Moreover, given a possible correlation between litter traits of the leaves and roots, we examined whether soil mesofauna alters the relationship between leaf and root decomposition across species. We found that the effects of plant species identity was stronger than that of soil mesofauna for determining the litter mass loss rate and the microbial respiration rate in both above‐ground and below‐ground decomposition. In addition, we found a significant positive correlation between leaf and root litter decomposition processes, regardless of the involvement soil mesofauna. On the other hand, the presence of soil mesofauna increased microbial respiration rates in the early stage of leaf decomposition; however, soil mesofauna did not affect root microbial respiration rates during the experiment. Such differential involvement of mesofauna in the leaf and root litter decomposition may drive the general patterns of faster and slower decomposition of plant leaves and roots in the soil, respectively.     

Litter mixture effects on tropical tree seedling growth--a greenhouse experiment

Decomposing litter provides critical nutrients for plants, particularly in nutrient-poor ecosystems such as tropical forests. We hypothesised that decomposing litter improves the performance of a variety of tropical tree seedlings, and that this litter effect varies depending on the species of litter present in litter mixtures. We addressed these hypotheses with a large pot experiment manipulating a range of different litter mixtures of contrasting quality and using seedlings of four tree species from the Amazonian forest of French Guiana. In contrast to our initial hypothesis, decomposing litter had either neutral or negative impacts on seedling growth, despite strongly different growth rates, biomass allocation patterns and leaf and root traits among tree species. Tree species varied in their responses to litter additions, which were further modified by species identity of the added litter. Our data show litter species-specific effects on growth, biomass allocation and leaf and root traits of tropical tree seedlings. These results suggest that a net nutrient release from decomposing litter does not necessarily improve tree seedling growth, even under nutrient-limiting conditions. In conclusion, litter layer composition may affect seedling establishment and recruitment success beyond litter-derived plant nutrient availability, which may contribute to tree species composition and dynamics in the studied tropical forest.     

Nutrient supply of forest soils in relation to management and site history

Internal and external factors like forest management practices and atmospheric deposition may have large influences on the nutrient supply of forest soils. Examples are given for the effects of tree species, harvesting, site history, changes of species and specific soil conditions, nitrogen deposition, and forest growth dynamics. It is concluded from these examples that all of these factors may contribute to soil acidification in forest ecosystems under humid temperate climatic conditions.     

Functional Composition of Tree Communities Changed Topsoil Properties in an Old Experimental Tropical Plantation

Forest biogeochemistry is strongly determined by the interaction between the tree community and the topsoil. Functional strategies of tree species are coupled to specific chemical leaf traits, and thus also to litter composition, which affects mineral soil characteristics. The limited understanding on this interaction is mainly based on shorter-term common garden experiments in temperate forest, and needs to be extended to other forest types and climates if we want to understand the universality of this linkage. In particular, for highly diverse tropical forests, our understanding of this interaction remains limited. Using an old experimental plantation within the central Congo basin, we examined the relationship between leaf and litter chemical composition and topsoil properties. Canopy, litter and topsoil characteristics were measured and we determined how the community-level leaf and litter chemical composition altered the topsoil carbon, major plant nutrients and exchangeable cation concentration, acidity and pH over the last eight decades. We found that functional composition strongly affected topsoil pH. In turn, topsoil pH strongly determined the soil total carbon and available phosphorus, total nitrogen and exchangeable potassium. Our results indicate that, as observed in temperate common garden experiments, trees alter chemical topsoil properties primarily through soil acidification, differently induced by functional composition of the tree community. The strong link between this community-level composition and topsoil characteristics, on a highly representative soil type for the tropics, improves our understanding of tropical forests biogeochemistry.     

Leaf litter stoichiometry affects decomposition rates and nutrient dynamics in tropical forests under restoration in Costa Rica

Active restoration strategies increase the production of leaf litter in tropical forests, but little is known about their effect on litter decomposition and subsequent nutrient release. We quantified changes in leaf litter stoichiometry during decomposition in former pasture sites under contrasting restoration strategies (natural regeneration, applied nucleation/islands tree planting and plantation), as well as in nearby primary forest. Litterbags were employed to evaluate decomposition. We used a leaf mixture of either the four planted tree species in the plantation and island treatments or the nearby primary forest and compared them under a factorial design. Decomposition rates were similar between restoration treatments (p > 0.5), but leaves decomposed faster in the forest mixture than in the plantation mixture (p < 0.01). The content of Ca, Mg, K, P, and the C:N ratio were higher in the forest mixture at the beginning and during decomposition (p < 0.05); the N content in the plantation mixture was higher at the beginning but lower during decomposition (p < 0.05), which meant greater mobilization of nitrogen per unit of carbon lost. K and P had a strong initial release, while Mg was released more gradually. N and Ca had an irregular pattern of initial fast release, immobilization, and re‐release in the later stages. We conclude that the differences in rates of decomposition and nutrient release in these systems under restoration were at least partly determined by the floristic heterogeneity and chemical quality of the leaf litter that reaches the soil.     

Influences of evergreen gymnosperm and deciduous angiosperm tree species on the functioning of temperate and boreal forests

It has been recognized for a long time that the overstorey composition of a forest partly determines its biological and physical–chemical functioning. Here, we review evidence of the influence of evergreen gymnosperm (EG) tree species and deciduous angiosperm (DA) tree species on the water balance, physical–chemical soil properties and biogeochemical cycling of carbon and nutrients. We used scientific publications based on experimental designs where all species grew on the same parent material and initial soil, and were similar in stage of stand development, former land use and current management. We present the current state of the art, define knowledge gaps, and briefly discuss how selection of tree species can be used to mitigate pollution or enhance accumulation of stable organic carbon in the soil. The presence of EGs generally induces a lower rate of precipitation input into the soil than DAs, resulting in drier soil conditions and lower water discharge. Soil temperature is generally not different, or slightly lower, under an EG canopy compared to a DA canopy. Chemical properties, such as soil pH, can also be significantly modified by taxonomic groups of tree species. Biomass production is usually similar or lower in DA stands than in stands of EGs. Aboveground production of dead organic matter appears to be of the same order of magnitude between tree species groups growing on the same site. Some DAs induce more rapid decomposition of litter than EGs because of the chemical properties of their tissues, higher soil moisture and favourable conditions for earthworms. Forest floors consequently tend to be thicker in EG forests compared to DA forests. Many factors, such as litter lignin content, influence litter decomposition and it is difficult to identify specific litter‐quality parameters that distinguish litter decomposition rates of EGs from DAs. Although it has been suggested that DAs can result in higher accumulation of soil carbon stocks, evidence from field studies does not show any obvious trend. Further research is required to clarify if accumulation of carbon in soils (i.e. forest floor + mineral soil) is different between the two types of trees. Production of belowground dead organic matter appears to be of similar magnitude in DA and EG forests, and root decomposition rate lower under EGs than DAs. However there are some discrepancies and still are insufficient data about belowground pools and processes that require further research. Relatively larger amounts of nutrients enter the soil–plant biogeochemical cycle under the influence of EGs than DAs, but recycling of nutrients appears to be slightly enhanced by DAs. Understanding the mechanisms underlying forest ecosystem functioning is essential to predicting the consequences of the expected tree species migration under global change. This knowledge can also be used as a mitigation tool regarding carbon sequestration or management of surface waters because the type of tree species affects forest growth, carbon, water and nutrient cycling.     

降水变化和氮沉降影响森林叶根凋落物分解研究进展

全球环境变化通过改变凋落物质量和产量、土壤生物以及非生物因子调控森林凋落物分解，从而对森林生态系统物质和能量循环产生重要的影响。就森林凋落物分解对当前我国面临降水格局变化和大气氮沉降增加的响应进行了回顾和系统的分析，发现降水格局改变如降水减少可能降低凋落物质量从而减缓凋落物分解，而氮沉降增加通常提高凋落物质量从而促进凋落物分解（间接效应）；降水格局改变通过调节土壤含水量和溶解氧含量进而影响微生物参与的分解过程，或通过改变可溶性组分的淋溶量来影响凋落物分解的物理过程，而氮沉降增加主要通过提高外源氮素的有效性从而促进或抑制微生物参与的分解过程（直接效应）。现有研究大多是基于地上凋落物（例如叶凋落物）来理解和量化森林凋落物分解速率与环境因子之间的关系。但目前对降水格局变化及其与大气氮沉降增加的交互作用如何影响森林地上和地下凋落物分解，以及潜在的微生物学机制仍然缺乏统一和清晰的认识。从土壤性质、凋落物质量、微生物群落结构和功能3个方面构建了环境变化对森林地上和地下凋落物分解的概念框架，并进一步阐述未来研究的重点方向：（1）亟需查明地上和地下凋落物分解的驱动机制；（2）探明降水格局变化和氮添加单因子及两因子交互作用对凋落物分解和养分释放的影响及其生物化学调控机理；（3）阐明微生物群落结构和功能对降水格局变化和氮添加单因子及两因子交互的响应机制。以期为深入探讨全球环境变化对森林凋落物分解的影响，以及环境胁迫下森林土壤"碳库"维持机制的解释提供科学依据。     

Partitioning the effect of composition and diversity of tree communities on leaf litter decomposition and soil respiration

The decomposition of plant material is an important ecosystem process influencing both carbon cycling and soil nutrient availability. Quantifying how plant diversity affects decomposition is thus crucial for predicting the effect of the global decline in plant diversity on ecosystem functioning. Plant diversity could affect the decomposition process both directly through the diversity of the litter, and/or indirectly through the diversity of the host plant community and its affect on the decomposition environment. Using a biodiversity experiment with trees in which both functional and taxonomic diversity were explicitly manipulated independently, we tested the effects of the functional diversity and identity of the living trees separately and in combination with the functional diversity and identity of the decomposing litter on rates of litter decomposition and soil respiration. Plant traits, predominantly leaf chemical and physical traits, were correlated with both litter decomposition and soil respiration rates. Surface litter decomposition, quantified by mass loss in litterbags, was best explained by abundance‐weighted mean trait values of tree species from which the litter was assembled (functional identity). In contrast, soil respiration, which includes decomposition of dissolved organic carbon and root respiration, was best explained by the variance in trait values of the host trees (functional diversity). This research provides insight into the effect of loss of tree diversity in forests on soil processes. Such understanding is essential to predicting changes in the global carbon budget brought on by biodiversity loss.     

Do physical plant litter traits explain non‐additivity in litter mixtures? A test of the improved microenvironmental conditions theory

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.     

Foliage litter turnover and earthworm populations in three beech forests of contrasting soil and vegetation types

Summary Leaf litter decomposition, levels of accumulated litter as well as the abundance and biomass of earthworms were measured in three mature beech forests in southern Sweden: one mor site, one poor mull site, and one rich mull site. The disappearance rate of beech litter, measured with litter bags, increased with increasing soil fertility. On the rich mull site, the disappearance rate was much higher than in the two other forests, due to the combined effects of higher earthworm activity, more favouable soil moisture conditions, and higher litter quality. Incubating the litter in finely meshed bags (1-mm mesh) to exclude macrofauna had a great effect on litter mass loss in the rich mull site, but it had only a minor effect in the other sites. Simultaneous incubations of local and transplanted leaf litter on the three study sites showed that the substrate quality of the litter increased in the order: mor site — poor mull site — rich mull site. Lignin, N, and P concentrations of the leaf litter failed to explain the observed differences in decomposition rates, and acid/base properties are suggested to be more important. Earthworm numbers per m² were 2.5 (1 species) in the mor, 40 (6 species) in the poor mull and 220 (9 species) in the rich mull forest. Soil chemical conditions, notably pH, were suggested as the main factors determining the inter-site differences in abundance and species composition of earthworms. The role of litter decomposition and earthworm activity in the accumulation of organic matter in the forest floor in different types of beech woodlands are discussed.     

Establishment of mixed plantations of Pinus sylvestris var. mongolica and Populus × xiaozhuanica may not be appropriate: evidence from litter decomposition

Aims Mongolian pine (Pinus sylvestrisvar.mongolica) and Xiaozhuan poplar (Populus × xiaozhuanica) are two predominant afforestation tree species in the semi-arid sandy lands of northeast China, which are characterized by poor soil nutrients. Plant litter decomposition plays a critical role in regulating nutrient cycling in terrestrial ecosystems. Admixture of broadleaf litter to conifer litter is expected to improve litter decomposition and soil fertility, and thus productivity. However, the effects on the decomposition of litter mixture of the above two tree species are not well understood. Therefore, it is essential to assess the decomposition performance of litter mixture with the aim of improving forest nutrient management and the establishment of mixed plantation. Appropriate forest management practice is critical for the sustainability of site productivity in plantation forests.     

长白山阔叶红松林主要树种凋落叶分解速率及其与叶性状的关系

阔叶红松林是我国东北地区地带性顶级森林群落,对维持区域生态系统稳定性具有重要作用。对阔叶红松林内主要树种凋落叶分解过程及影响因素进行研究,有助于增加长白山阔叶红松林生态系统的基础数据,为明确阔叶红松林的养分循环和物质流动提供依据。选取了长白山阔叶红松林内30个常见乔灌树种和16个凋落叶性状,采用野外分解袋法和室内样品分析等方法研究了长白山阔叶红松林内主要树种凋落叶分解速率及其与凋落叶性状的关系。1年的野外分解实验表明,30个树种的凋落叶重量损失率表现出较大差异。不同树种凋落叶的重量损失率在20.56%—92.11%之间,以红松(Pinus koraiensis)质量损失率最低,东北山梅花(Philadelphus schrenkii)质量损失率最高。不同生活型树种的凋落叶在质量损失率上存在显著差异,以灌木树种凋落叶的质量损失率最高,小乔木次之,乔木树种质量损失率最低。Olson模型拟合结果表明,不同树种凋落叶的分解速率k以红松最低,瘤枝卫矛(Euonymus verrucosus)最高,分别为0.24和1.64。不同树种分解50%和95%所需的时间分别在0.43—2.86年,1.83—...     

Nitrogen Addition Shapes Soil Phosphorus Availability in Two Reforested Tropical Forests in Southern China

Scant information is available on how soil phosphorus (P) availability responds to atmospheric nitrogen (N) deposition, especially in the tropical zones. This study examined the effect of N addition on soil P availability, and compared this effect between forest sites of contrasting land‐use history. Effects of N addition on soil properties, litterfall production, P release from decomposing litter, and soil P availability were studied in a disturbed (reforested pine forest with previous understory vegetation and litter harvesting) and a rehabilitated (reforested mixed pine/broadleaf forest with no understory vegetation and litter harvesting) tropical forest in southern China. Experimental N‐treatments (above ambient) were the following: Control (no N addition), N50 (50 kg N ha^?1 yr^?1), and N100 (100 kg N ha^?1 yr^?1). Results indicated that N addition significantly decreased soil P availability in the disturbed forest. In the rehabilitated forest, however, soil P availability was significantly increased by N addition. Decreases in soil P availability may be correlated with decreases in rates of P release from decomposing litter in the N‐treated plots, whereas the increase in soil P availability was correlated with an increase in litterfall production. Our results suggest that response of soil P availability to N deposition in the reforested tropical forests in southern China may vary greatly with temporal changes in tree species composition and soil nutrient status, caused by different land‐use practices.     

Species-specific characteristics of trees can determine the litter macroinvertebrate community and decomposition process below their canopies

This paper tests whether individual trees in a mature forest stand influence the process of litter decomposition and the macroinvertebrate communities in the soil underneath their canopies, as a result of species-specific characteristics. A field decomposition experiment was performed in a mature forest stand of tropical montane cloud forest in Mexico. The areas under the canopies of Quercus laurina Humbl. & Bompl., Oreopanax xalapensis (Kunth) Decne. & Planchon and Beilschmedia ovalis (Blake) C. K. Allen trees were used as experimental units. The natural soil and litter macroinvertebrate communities were monitored and compared to the community that invaded decomposition boxes with reciprocally transplanted leaf litter. The abundances of four macroinvertebrate taxa in natural litter differed among tree species independently of season. No differences were found in the soil community. The response to experimental litter by macroinvertebrate taxa suggests that the production of a specific quality of litter is an important mechanism by which a tree influences the litter macroinvertebrate community that develops under its canopy. However, not all differences in community composition naturally found between tree species can be explained by differences in litter quality during the first year of decomposition. Differences in nutrient release that occur after the first year, and physical properties of litter also probably play an important role. Independently of the canopy tree, the initial chemical quality (N, P, Ca, Mg and lignin) of experimental litter largely determined the decomposition rate and nutrient dynamics of decomposing leaves. However, it was found that under O. xalapensis trees the breakdown of lignin from the litter produced by the same species of tree was particularly effective. This suggests that a feedback has developed between this tree species and the decomposer community prevailing under its canopy.