Litter dynamics recover faster than arthropod biodiversity during tropical forest succession

Litterfall and litter decomposition are key elements of nutrient cycling in tropical forests, a process in which decomposer communities such as macro-arthropods play a critical role. Understanding the rate and extent to which ecosystem function and biodiversity recover during succession is useful to managing the growing area of tropical successional forest globally. Using a replicated chronosequence of forest succession (5–15, 15–30, 30–45 years, and primary forest) on abandoned pastures in lowland tropical wet forest, we examined litterfall, litter chemistry, and effects of macro-arthropod exclusion on decomposition of two litter types (primary and 5- to 15-years-old secondary forest). Further, we assessed macro-arthropod diversity and community composition across the chronosequence. Overstory cover, litterfall, and litter nutrients reached levels similar to primary forest within 15–30 years. Young secondary forest litter (5–15 years) had lower initial N and P content, higher C:N, and decayed 60 percent faster than primary forest litter. The presence of macro-arthropods strongly mediated decomposition and nutrient release rates, increasing litter mass loss by 35–44 percent, N released by 53 percent, and P release by 84 percent. Forest age had no effect on soil nutrients, rates of litter decomposition, nutrient release, or macro-arthropod influence. In contrast, abundance and community composition of macro-arthropods remained significantly lower and distinct in all ages of secondary compared with primary forest. Order richness was lower in 5–15 years of secondary compared with primary forest. Our results suggest that in highly productive tropical wet forest, functional recovery of litter dynamics precedes recovery of decomposer community structure and biodiversity.     

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.     

季节性雪被变化对森林凋落物分解及土壤氮动态的影响

全球气候变化引发的雪被格局变化将深刻影响植被的凋落物分解、陆地生态系统的土壤养分循环等过程.森林是陆地生态系统的主体,在全球生物地球化学循环中起着不可替代的作用.本研究综述了季节性雪被变化对森林凋落物分解及土壤氮动态的影响.全球气候变化情景下季节性雪被表现出因地域而异的增加或减少的变化格局,一方面通过改变环境温湿度、凋落物质量、分解者动态等直接影响分解过程,另一方面通过改变森林群落结构、植被物候、土壤养分等间接地作用于凋落物分解.同时,季节性雪被通过影响氮富集作用、雪被下土壤温湿度、冻融循环、森林群落、雪下动物和微生物等相关因子而改变森林土壤氮循环.本领域未来应开展的研究是: 1) 全面考虑全球气候变化情景下季节性雪被格局的变异性,开展不同季节性雪被格局变化的模拟研究;2) 开展季节性雪被融雪水淋溶作用对森林凋落物分解和土壤氮动态的影响研究;3) 阐明不同生态系统和气候带中季节性雪被格局变化对森林凋落物分解过程和土壤氮动态的驱动机制研究;4) 量化季节性雪被变化对森林凋落物分解和土壤氮动态在雪被覆盖期的瞬时影响和无雪期的延续影响,为阐明和模型预测陆地生态系统生物地球化学循环对全球气候变化的响应提供理论基础和数据支持.     

Nutrient fluxes via litterfall and leaf litter decomposition vary across a gradient of soil nutrient supply in a lowland tropical rain forest

The extent to which plant communities are determined by resource availability is a central theme in ecosystem science, but patterns of small-scale variation in resource availability are poorly known. Studies of carbon (C) and nutrient cycling provide insights into factors limiting tree growth and forest productivity. To investigate rates of tropical forest litter production and decomposition in relation to nutrient availability and topography in the absence of confounding large-scale variation in climate and altitude we quantified nutrient fluxes via litterfall and leaf litter decomposition within three distinct floristic associations of tropical rain forest growing along a soil fertility gradient at the Sepilok Forest Reserve (SFR), Sabah, Malaysia. The quantity and nutrient content of small litter decreased along a gradient of soil nutrient availability from alluvial forest (most fertile) through sandstone forest to heath forest (least fertile). Temporal variation in litterfall was greatest in the sandstone forest, where the amount of litter was correlated negatively with rainfall in the previous month. Mass loss and N and P release were fastest from alluvial forest litter, and slowest from heath forest litter. All litter types decomposed most rapidly in the alluvial forest. Stand-level N and P use efficiencies (ratios of litter dry mass to nutrient content) were greatest for the heath forest followed by the sandstone ridge, sandstone valley and alluvial forests, respectively. We conclude that nutrient supply limits productivity most in the heath forest and least in the alluvial forest. Nutrient supply limited productivity in sandstone forest, especially on ridge and hill top sites where nutrient limitation may be exacerbated by reduced rates of litter decomposition during dry periods. The fluxes of N and P varied significantly between the different floristic communities at SFR and these differences may contribute to small-scale variation in species composition.     

Stronger effects of litter origin on the processing of conifer than broadleaf leaves: A test of home‐field advantage of stream litter breakdown

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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。     

Comparison of Leaf Litter Decomposition Rates in Restored and Mature Riparian Forests on the Sacramento River, California

One of the largest riparian restoration projects in the United States is currently taking place in California on the Sacramento River. Nearly 2,000 ha of land adjacent to the river channel have been revegetated with native riparian species in an effort to reestablish riparian forests. The objective of this study was to compare leaf litter decomposition rates in restored riparian forests to those in mature, naturally established riparian forests, in order to monitor the development of this ecosystem function in restored forests. Leaf litter decomposition rates were measured over 1 year in six restored riparian forests (4, 7, and 9 years old) and two mature remnant riparian forests (>50 years old), in order to test two hypotheses: (1) decomposition rates of restored and mature forests are significantly different and (2) decomposition rates in the chronosequence of restored forests are moving along a trajectory, approaching the decomposition rates characteristic of mature forests as they age. Statistical analyses revealed no significant differences in annual decay rates among the four different forest ages and no trajectory among leaf litter decomposition rates in restored forests. These results suggest that a functionally equivalent process of leaf litter decomposition occurs in both restored and naturally established forests and show promise for the efficiency of nutrient cycling processes in these restored forests.     

All size classes of soil fauna and litter quality control the acceleration of litter decay in its home environment

There is increasing evidence that litter decomposition is faster beneath the plant species it was derived from, an effect called home‐field advantage (HFA). Adaptation of soil biota to decompose the litter that they encounter most often has been proposed as the main mechanism to explain HFA. However, there is little direct evidence supporting this assumption and the contribution of different decomposer groups to the HFA is unknown. Furthermore, the large observed variation in the strength of the HFA may be linked to litter quality, with increasing HFA for more recalcitrant substrates. To test the relationship between HFA, litter quality and soil fauna, we performed a field litter bag experiment, with different mesh sizes and reciprocal litter transplants, across a successional gradient varying in litter quality inputs and soil fauna composition. The results show that the HFA was stronger in sites dominated by more recalcitrant litter inputs and provide evidence that a range of soil fauna size classes contribute to this effect. Our findings help explain the lack of HFA in ecosystems with simple litters and in studies where experimental artefacts (e.g. fine mesh sizes) affect the contribution from certain classes of soil fauna.     

Spatial variation in leaf nutrient traits of dominant desert riparian plant species in an arid inland river basin of China

Understanding how patterns of leaf nutrient traits respond to groundwater depth is crucial for modeling the nutrient cycling of desert riparian ecosystems and forecasting the responses of ecosystems to global changes. In this study, we measured leaf nutrients along a transect across a groundwater depth gradient in the downstream Heihe River to explore the response of leaf nutrient traits to groundwater depth and soil properties. We found that leaf nutrient traits of dominant species showed different responses to groundwater depth gradient. Leaf C, leaf N, leaf P, and leaf K decreased significantly with groundwater depth, whereas patterns of leaf C/N and leaf N/P followed quadratic relationships with groundwater depth. Meanwhile, leaf C/P did not vary significantly along the groundwater depth gradient. Variations in leaf nutrient traits were associated with soil properties (e.g., soil bulk density, soil pH). Groundwater depth and soil pH jointly regulated the variation of leaf nutrient traits; however, groundwater depth explained the variation of leaf nutrient traits better than did soil pH. At the local scale in the typical desert riparian ecosystem, the dominant species was characterized by low leaf C, leaf N, and leaf P, but high leaf N/P and leaf C/P, indicating that desert riparian plants might be more limited by P than N in the growing season. Our observations will help to reveal specific adaptation patterns in relation to the groundwater depth gradient for dominant desert riparian species, provide insights into adaptive trends of leaf nutrient traits, and add information relevant to understanding the adaptive strategies of desert riparian forest vegetation to moisture gradients.     

Ectomycorrhizal-Dominated Boreal and Tropical Forests Have Distinct Fungal Communities,but Analogous Spatial Patterns across Soil Horizons

Fungi regulate key nutrient cycling processes in many forest ecosystems, but their diversity and distribution within and across ecosystems are poorly understood. Here, we examine the spatial distribution of fungi across a boreal and tropical ecosystem, focusing on ectomycorrhizal fungi. We analyzed fungal community composition across litter (organic horizons) and underlying soil horizons (0–20 cm) using 454 pyrosequencing and clone library sequencing. In both forests, we found significant clustering of fungal communities by site and soil horizons with analogous patterns detected by both sequencing technologies. Free-living saprotrophic fungi dominated the recently-shed leaf litter and ectomycorrhizal fungi dominated the underlying soil horizons. This vertical pattern of fungal segregation has also been found in temperate and European boreal forests, suggesting that these results apply broadly to ectomycorrhizal-dominated systems, including tropical rain forests. Since ectomycorrhizal and free-living saprotrophic fungi have different influences on soil carbon and nitrogen dynamics, information on the spatial distribution of these functional groups will improve our understanding of forest nutrient cycling.     

喀斯特峰丛洼地不同类型森林养分循环特征

以中国西南喀斯特峰丛洼地为研究区域用标准木法和收获法对人工林、次生林、原生林3个不同类型森林的6个代表性群落的生物量、营养元素生物循环量及循环特征进行了研究。结果表明:(1)不同类型森林群落乔木各器官的养分含量大小顺序为:叶枝根干,林下植被层和凋落物层的养分含量比较高,其含量普遍高于乔木层各组分,仅次于乔木叶片;各组分中营养元素以K、Ca最高,P、Mg最低;(2)3种类型森林间乔木层的养分积累量总规律表现为原生林(4540.30 kg/hm~2)次生林(2107.09 kg/hm~2)人工林(719.51 kg/hm~2),分别占林分养分积累量的88.30%、79.57%和62.60%;(3)3种类型森林生态系统养分总贮量相差不大,均主要集中在土壤层在各层分配格局有所差异;营养元素的年吸收量和年归还量均为次生林原生林人工林,年吸收量分别为:418.80、271.17和148.79 kg hm~(-2)a~(-1);年归还量分别为:182.98、111.43和43.37 kg hm_(-2)a~(-1);(4)不同类型森林养分利用系数总规律为人工林(0.35)次生林(0.20)原生林(0.10);循环系数则相反,为原生林(0.48)次生林(0.46)人工林(0.30);而周转时间为原生林(37.32)人工林(18.63)次生林(13.93)。喀斯特峰丛洼地土层薄,养分贮存能力差,森林养分循环能力相对较弱,沿着强、中、弱干扰递减梯度,3种类型森林养分利用效率和循环能力呈增长趋势。     

A Meta-analysis of Climatic and Chemical Controls on Leaf Litter Decay Rates in Tropical Forests

Although tropical forests occupy a small fraction of the earth’s total land area, they play a disproportionately large role in regulating the global carbon cycle. Yet controls on both primary productivity and decomposition in tropical forests are not well-studied in comparison with temperate forests and grasslands, despite their extreme biogeochemical heterogeneity. To evaluate the relative importance of climate and foliar chemical variables in driving decomposition in tropical forests, I performed a meta-analysis of reported leaf litter decay rates throughout tropical forest ecosystems. Using a model selection procedure based on Akaike’s Information Criterion, I found that temperature and precipitation played little direct role in regulating decomposition rates, except in montane forests where cool temperatures slowed decay. Foliar concentrations of calcium, magnesium, nitrogen, phosphorus, and potassium were important predictors of mass loss rates, although each of these factors explained a very small amount of variance when considered in isolation. The large amount of unexplained variation in decomposition rates observed both within and across tropical forest sites may be due to other factors not explored here, such as soil biota or complex plant secondary chemistry. Carbon cycling in tropical forests seems to be modulated by the availability of multiple nutrients, underscoring the need for additional manipulative experiments to explore patterns of belowground nutrient limitation across the biome. Because models of decomposition developed in temperate ecosystems do not appear to be generalizable to wet tropical forests, new biogeochemical paradigms should be developed to accommodate their unique combination of climatic, edaphic, and biotic factors.     

Defining nutritional constraints on carbon cycling in boreal forests – towards a less `phytocentric' perspective

Growing interest in possible global climate change has underlined the need for better information concerning the way in which carbon partitioning between ecosystem components is influenced by constraints on nutrient availability. Micro-organisms play a fundamental role in the cycling of carbon and nutrients in all ecosystems but the role of fungi in particular is pivotal in boreal forest ecosystems. Traditional models of nutrient cycling are based on methods and concepts developed in agricultural systems where microorganisms are considered primarily as nutrient processors providing plants with inorganic nutrients. The filamentous nature of fungi, their ability to translocate carbon and nutrients between different substrates and the capacity of ectomycorrhizal fungi to utilise organic nutrients have all been largely ignored. In this article, a new model is suggested which emphasises competition for organic nutrients between decomposer organisms and plants, with the plants depending on their associated mycorrhizal fungi for nutrient acquisition. Antagonistic interactions involving nutrient transfer between decomposer and mycorrhizal fungi are proposed as important pathways in nutrient cycling. Due to the nutrient conservative features of decomposer fungi, inorganic nutrients are considered less important for plant nutrition. The implications of the new nutrient cycling model on the carbon balance of boreal forests are discussed.     

Using experimental manipulation to assess the roles of leaf litter in the functioning of forest ecosystems

The widespread use of forest litter as animal bedding in central Europe for many centuries gave rise to the first litter manipulation studies, and their results demonstrated that litter and its decomposition are a vital part of ecosystem function. Litter plays two major roles in forest ecosystems: firstly, litterfall is an inherent part of nutrient and carbon cycling, and secondly, litter forms a protective layer on the soil surface that also regulates microclimatic conditions. By reviewing 152 years of litter manipulation experiments, I show that the effects of manipulating litter stem from changes in one, or both, of these two functions, and interactions between the variables influenced by the accumulation of litter can result in feedback mechanisms that may intensify treatment effects or mask responses, making the interpretation of results difficult.Long-term litter removal increased soil bulk density, overland flow, erosion, and temperature fluctuations and upset the soil water balance, causing lower soil water content during dry periods. Soil pH increased or decreased in response to manipulation treatments depending on forest type and initial soil pH, but it is unclear why there was no uniform response. Long-term litter harvesting severely depleted the forests of nutrients. Decreases in the concentrations of available P, Ca, Mg, and K in the soil occurred after only three to five years. The decline in soil N occurred over longer periods of time, and the relative loss was greater in soils with high initial nitrogen concentration. Tree growth declined with long-term litter removal, probably due to lower nutrient availability. Litter manipulation also added or removed large amounts of carbon thereby affecting microbial communities and altering soil respiration rates.Litter manipulation experiments have shown that litter cover acts as a physical barrier to the shoot emergence of small-seeded species; further, the microclimate maintained by the litter layer may be favourable to herbivores and pathogens and is important in determining later seedling survival and performance. Litter manipulation altered the competitive outcomes between tree seedlings and forbs, thereby influencing species composition and diversity; changes in the species composition of understorey vegetation following treatments occurred fairly rapidly. By decreasing substrate availability and altering the microclimate, litter removal changed fungal species composition and diversity and led to a decline in populations of soil fauna. However, litter addition did not provoke a corresponding increase in the abundance or diversity of fungi or soil fauna.Large-scale long-term studies are still needed in order to investigate the interactions between the many variables affected by litter, especially in tropical and boreal forests, which have received little attention. Litter manipulation treatments present an opportunity to assess the effects of increasing primary production in forest ecosystems; specific research aims include assessing the effects of changes in litter inputs on the carbon and nutrient cycles, decomposition processes, and the turnover of organic matter.     

The legacy of forest logging on organic matter inputs and storage in tropical streams

Riparian forests play an important role in stream ecosystems, as they support biodiversity, reduce water erosion, and provide litter that fuels aquatic biota. However, they are affected by great array of anthropogenic threats (e.g., fire, logging, and organic pollution), which alter species composition and their physical structure. Although forest recovery after disturbance such as logging can take decades, the legacy of forest clear-cut logging on key processes in tropical riparian ecosystems is mostly unknown. Here, we investigated how litter inputs (leaves, twigs, and reproductive parts) and storage, key processes for carbon and nutrient recycling and for forest and stream biota, are influenced by riparian vegetation undergoing succession (after 28 years from logging) through the comparison of reference and logged forest sites in the Cerrado biome. Litterfall was overall similar between forest types, but litterfall of twigs was twofold higher at logged than reference sites. Similarly, litter inputs from the bank to the stream (i.e., lateral inputs) and streambed storage were 50–60% higher at logged than reference sites. The higher litterfall observed in logged forests could be related to higher proportion of tree species that are characteristic of primary and secondary successional stages, including fast-growing and liana species, which often are more productive and common in anthropogenic areas. Our results showed that the legacy impact of clear-cut logging, even if residual woody vegetation is maintained in riparian buffers, can shift the type, quantity, and seasonality of litter subsidies to tropical streams. This knowledge should be considered within the context of management and conservation of communities and ecosystem processes in the forest-stream interfaces.     

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.     

Variable Responses of Lowland Tropical Forest Nutrient Status to Fertilization and Litter Manipulation

Predicting future impacts of anthropogenic change on tropical forests requires a clear understanding of nutrient constraints on productivity. We compared experimental fertilization and litter manipulation treatments in an old-growth lowland tropical forest to distinguish between the effects of inorganic nutrient amendments and changes in nutrient cycling via litterfall. We measured the changes in soil and litter nutrient pools, litterfall, and fine root biomass in plots fertilized with nitrogen (N), phosphorus (P), or potassium (K), and in litter addition and litter removal treatments during 7 years. Soil inorganic N and litter N increased in double-litter plots but not in N-fertilized plots. Conversely, litter P and soil pools of P and K increased in fertilized plots but not in the double-litter plots. Soil and litter pools of N and K decreased in the no-litter plots. Changes in litterfall with added nutrients or litter were only marginally significant, but fine root biomass decreased with both the litter and the K addition. Differences between the two experiments are mostly attributable to the coupled cycling of carbon and nutrients in litter. Increased nutrient inputs in litter may improve plant uptake of some nutrients compared to fertilization with similar amounts. The litter layer also appears to play a key role in nutrient retention. We discuss our findings in the context of possible impacts of anthropogenic change on tropical forests.     

内蒙古东部天然白桦林的凋落物性质和储量及其随温度和降水梯度的变化格局

作为碳素和养分循环的重要组分, 地表凋落物如何响应全球气候变化日益受到重视。中国北方半干旱地区森林草原过渡带斑块状分布的森林对气候变化引起的水热变化的响应较为敏感, 但是对这些森林地表处于不同分解阶段的凋落物的化学性质和储量的格局及其如何响应温度、降水变化的报道较少。该研究分析了内蒙古东部地区处于年平均气温和年降水量梯度上的12个天然白桦(Betula platyphylla)林不同凋落物层次(最上层的初步分解层L1, 中间的半分解层L2, 最下层的腐殖质层L3)的化学性质及现存量后发现: 1)随着分解的进行(即从L1到L3层), 氮、磷浓度显著增加, 可提取物浓度基本不变, 酸溶性组分(acid soluble fraction, AS)浓度下降, 酸不溶性组分(acid insoluble fraction, AIF)浓度增加。2)各元素现存量均在L3层最高, 表明凋落物分解缓慢, 养分积累。3)年平均气温和年降水量对凋落物有机组分(AS组分和AIF组分)的性质无显著影响, 但L3层元素储量随年平均气温升高而增加, 可能由于年平均气温较高的地点森林生产力更高, 从而导致叶凋落物量增加, 但由于受水分限制(尤其是在夏天), 这些地点的凋落物分解速率不变或更低, 使凋落物积累更为明显。上述结果表明: 腐殖质层是这些白桦林的一个重要的碳及养分库, 未来在降水没有明显变化的情况下, 这一区域的升温可能会增加白桦林地表凋落物储量。     

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.