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.     

森林演替过程中优势树种凋落叶对土壤微生物组成的影响

为了进一步认识森林群落演替机理,研究了黄土高原马栏林区处于不同演替阶段的森林土壤微生物组成,以及优势树种凋落叶对土壤微生物组成的影响.采用凋落叶浸提液处理不同类型的土壤,利用土壤浸提液固体培养基测定微生物数量,以系统聚类和主成分分析等方法进行数据分析.结果表明:微生物总数量及土壤细菌数量按白桦(Betulaplatyphylla)林→油松(Pinus tabulaeformis)林→辽东栎(Quercus liaotungensis)林递增;在先锋森林群落向顶极群落演替过程中,微生物组成的综合性指标Shannon-Wiener指数呈下降趋势;土壤微生物3大类群中,数量较少者对Shannon-Wiener指数的大小变化起着较大作用;依据可培养微生物组成,8种土样可划分为3个类群,辽东栎及油松凋落叶对土壤微生物组成的影响不同,辽东栎凋落叶对土壤微生物学性质的作用更强;马栏林区森林土壤微生物组成与植被类型及演替阶段关系密切;演替高级阶段森林优势树种凋落叶可能通过其对土壤微生物组成的改造优势,逐渐改变土壤的微生物学性质,进而促进植被演替.     

泰山4种优势造林树种叶片凋落物分解对凋落物内细菌群落结构的影响

为研究泰山不同造林树种凋落物叶分解对细菌群落的影响。以泰山4种主要优势造林树种刺槐(Robinia pseucdoacacia)、麻栎(Quercus acutissima)、油松(Pinus tabulaeformis)和赤松(Pinus densiflora)为研究对象,采用凋落物分解袋法及Illumina Miseq测序平台对细菌16S rDNA V4—V5区扩增产物进行双端测序,分析了4种树种叶片凋落物分解对细菌群落结构及多样性的影响。结果表明:(1)4种树种叶片分解速率差异显著(P0.05),刺槐分解速率显著高于其他3个树种(P0.05),表现为刺槐赤松油松麻栎。(2)4种叶凋落物分解一年后化学元素含量与初始化学元素相比均存在显著差异。C、木质素含量均显著降低(P0.05);N、P含量显著升高(P0.05)。(3)所有样品一共获得643440条有效序列,分属于35门,92纲,121目,246科,410属,206种。细菌群落NMDSβ-多样性分析显示除油松和赤松间差异较小外,其他树种间差异程度均较大。其中,细菌群落相对丰度在5%以上的优势类群是变形菌门、放线菌门、拟杆菌门、酸杆菌门,且在4种处理之间差异显著(PSymbol|@@0.05)。在纲水平上,α-变形菌纲、β-变形菌纲、不明放线菌纲、鞘脂杆菌纲、γ-变形菌纲、δ-变形菌纲为主要的优势纲,其中不明放线菌纲和鞘脂杆菌纲差异显著(PSymbol|@@0.05)。在种水平上,Bradyrhizobium elkanii和Luteibacter rhizovicinus在4个处理中都为优势种,每个处理也都有自己所特有的优势种。(4)4个处理细菌丰富度(OUT、观测到的物种数和ACE指数)和系统发育多样性(PD指数)之间差异显著(PSymbol|@@0.05),且阔叶树种刺槐和麻栎显著高于针叶树种赤松和油松。(5)叶片凋落物性状和细菌群落NMDS分析表明,细菌群落多样性受到凋落物化学性质的影响,尤其是凋落物初始C/N比和木质素/N比。此外,在细菌群落多样性和叶片凋落物化学性质两个因素中,分解速率与凋落物化学性质相关性更大。研究结果有助于理解细菌群落结构和多样性对森林生态系统叶片凋落物分解的影响。     

Effects of leaf litter on woody seedlings in xeric successional communities

We investigated the effect of leaf litter on the establishment of Eucalyptus incrassata, a mallee eucalypt. It has been suggested that litter accumulation may hinder seedling establishment, and that the removal of litter may be one of the mechanisms through which fire enhances recruitment. We conducted factorial experiments testing the effects of three kinds of leaf litter on E. incrassata seeds and seedlings at three contiguous sites with different land use histories. One site was an uncleared E. incrassata open mallee woodland (Mallee site), one a cleared area that had been ungrazed for about five years (Pasture site) and the third an area of mallee rolled some 40 years ago and permitted to regenerate (Regrowth site). Litter had no effect on emergence of planted E. incrassata seeds, but emergence differed between sites. Overall, the percentage of seeds that germinated and emerged was substantial (mean 35.2% ± 25.9%). Seedling shoot biomass did not differ between sites or litter treatments. Although seedlings grown in Pasture litter suffered higher mortality rates, overall mortality rates were low (mean 13.2% ± 15.5%), suggesting that leaf litter has little effect on recruitment rates during winter and spring. We conclude that leaf litter does not affect emergence or growth in young E. incrassata seedlings during winter and spring, when most establishment occurs. Our results emphasize the difficulty in predicting litter effects on recruitment.     

The influence of variation in litter habitats on spider communities

Summary Spider communities were sampled over an artificial gradient of litter depth (created by raking) and compared to those of two other forests exhibiting natural variation in litter depth. More species of spiders were present in areas of greater depth and/or complexity in all sites. Relative abundance of Lycosidae decreased, while relative abundance of Clubionidae, Thomisidae and Gnaphosidae increased over gradients of increasing depth and complexity. Similarity of species composition between areas within a forest was related to site differences in litter depth and structure.As litter depth increased, there were significant changes in prey species richness, litter complexity, and microclimate. Partial correlation analysis of grouped data from early-, mid-, and late season suggests that influential factors change with season. In the early season, prey abundance and temperature variation account for most of the variation in spider species richness. In mid-season, litter complexity and moisture fluctuations appear to influence richness, with complexity relatively more important. In late season, complexity and temperature range were the primary factors, with temperature relatively more influential. The relative importance of these factors in influencing community structure of spiders is discussed.     

Variable effects of air-drying on leaching losses from tree leaf litter

Leaching of soluble substances may be an important first step in leaf litter decomposition in small streams, but recent research has suggested that large leaching losses (up to 30% of initial mass in 48 h) may be an artifact created by using air-dried leaves in decomposition experiments. In laboratory experiments, we compared 3 d leaching losses from freshly fallen and air-dried senescent leaves of 27 tree species from different regions across Canada. Air-dried leaves from all species leached measurable amounts of original mass (3.6–32.8% dry mass), but leaching losses from fresh leaves (0–35%) were detectable in all but two species. Air-drying increased leaching losses in many species, but in others it reduced leaching losses or had no measurable effect. Results for leaves of the same species collected in different regions or in different years were generally similar, but species within the same genus often behaved very differently. Neither moisture content (fresh or air-dried), leaf thickness, nor cuticle thickness proved of any value as predictors of leaching losses or the effect of air-drying. The propensity of autumn-fallen leaves to leach, whether fresh or air-dried, appears to be a property of the individual tree species.     

模拟增温对湿地植物凋落物分解及细菌群落的影响

植物凋落物分解是湿地生态系统物质循环的重要组成部分,随着全球气候变化的逐渐加剧,气候变暖对湿地植物凋落物分解的影响已引起人们的广泛关注。本研究通过凋落物袋法对比研究了山东省南四湖湿地中芦苇和香蒲两种典型湿地植物的凋落物分解过程,利用开顶式生长室(Open-top Chamber, OTC)模拟了大气增温(2.0±0.5)℃—(4.0±0.5)℃对凋落物分解特征和细菌群落的影响。结果显示,增温显著加速了两种植物凋落物的分解速率,而木质素/氮(Lignin/N)、纤维素/氮(Cellulose/N)是影响凋落物分解速率的重要因子,与分解速率呈显著负相关。增温显著增加了细菌群落的丰度和多样性,碳是厚壁菌门(Firmicutes)等细菌丰度变化的驱动因子,而木质素、木质素/氮是拟杆菌门(Bacteroidota)等细菌丰度变化的驱动因子。细菌群落共现网络显示,在增温条件下,凋落物分解的细菌群落网络主要由共生关系组成。气候变暖提高了细菌之间的相互关系和互惠程度,加快了植物凋落物的分解进程,进而影响了湿地生态系统的碳收支平衡。     

Soil microbial communities adapt to genetic variation in leaf litter inputs

Plant genotypes can have important community‐ and ecosystem‐level effects. However, whether the extended phenotypes of plants feed back to influence the fitness of causal genotypes through soil processes remains unknown. We investigated whether aspen genotypes create distinct soil microbial communities that could potentially affect plant fitness. Using naturally occurring aspen stands in an old‐field system, we set up reciprocal litter transplants among ten genetically distinct aspen clones and tracked decomposition and changes in belowground nutrients and microbial communities for three years. We found that belowground microbial communities became adapted to process specific genotypes of aspen litter to the extent allowable by environment and litter chemistry. Belowground processes were driven by a combination of little quality and prior exposure to specific genotypes of litter. In general, litter from aspen genotypes native to the soil community decomposed more rapidly than did litter from foreign aspen genotypes (i.e. a home‐field advantage existed). While home‐field advantages have been documented to occur among litters of different species, we show that intraspecific variation can elicit similar, albeit weak, effects within a single species. Because rapid decomposition and nutrient cycling is likely to benefit fast‐growing, early‐successional species such as aspen, genotype‐mediated selection for soil microbial communities may feed back to positively affect plant fitness. In addition, belowground communities exhibited significant shifts in response to leaf litter inputs. When exposed to foreign litter, microbial communities changed to become more similar to the microbial community beneath the foreign litter's origin, indicating that belowground microbial communities are predictable given the genotype of the aboveground aspen clone.     

The influence of sampling method on the classification of wetland macroinvertebrate communities

Macroinvertebrate communities sampled by a corer, plankton net and sweep net from five wetlands on the Swan Coastal Plain were compared. The composition of the fauna collected in sweeps and tows was generally similar and differed from that collected in the cores. Cores caught fewer species than tows and sweeps at all wetlands and did not capture fast swimming hemipterans or less abundant taxa. The highest species richness was recorded in sweep samples in four out of the five wetlands. Classification (TWIN-SPAN) and ordination (SSH) of the samples collected in sweeps and tows gave good separation of the wetlands, whereas classification of core samples did not. Coring appeared to be the least suitable sampling method for describing the major components of the macroinvertebrate communities of these wetlands. Plankton tows were useful if the time available for sorting was limited as these samples were free of sediments and generally gave similar results to those obtained with sweeps. Sweeps appeared to be the most useful method for a large classification study as they collected more species and resulted in the best discrimination amongst wetlands.     

Resource–consumer diversity: testing the effects of leaf litter species diversity on stream macroinvertebrate communities

1. Understanding relationships between resource and consumer diversity is essential to predicting how changes in resource diversity might affect several trophic levels and overall ecosystem functioning. 2. We tested for the effects of leaf litter species diversity (i.e. litter mixing) on litter mass remaining and macroinvertebrate communities (taxon diversity, abundance and biomass) during breakdown in a detritus‐based headwater stream (North Carolina, U.S.A.). We used full‐factorial analyses of single‐ and mixed‐species litter from dominant riparian tree species with distinct leaf chemistries [red maple (Acer rubrum), tulip poplar (Liriodendron tulipifera), chestnut oak (Quercus prinus) and rhododendron (Rhododendron maximum)] to test for additivity (single‐species litter presence/absence effects) and non‐additivity (emergent effects of litter species interactions). 3. Significant non‐additive effects of litter mixing on litter mass remaining were explained by species composition, but not richness, and litter‐mixing effects were variable throughout breakdown. Specifically, small differences in observed versus expected litter mass remaining were measured on day 14; whereas observed litter mass remaining in mixed‐species leaf packs was significantly higher on day 70 and lower on day 118 than expected from data for single‐species leaf packs. 4. Litter mixing had non‐additive effects on macroinvertebrate community structure. The number of species in litter mixtures (two to four), but not litter species composition, was a significant predictor of the dominance of particular macroinvertebrates (i.e. indicator taxa) within mixed‐species packs. 5. In addition, the presence/absence of high‐ (L. tulipifera) and low‐quality (R. maximum) litter had additive effects on macroinvertebrate taxon richness, abundance and biomass. The presence of L. tulipifera litter had both positive (synergistic) and negative (antagonistic) effects on invertebrate taxon richness, that varied during breakdown but were not related to litter chemistry. In contrast, the presence/absence of L. tulipifera had a negative relationship with total macroinvertebrate biomass (due to low leaf mass remaining when L. tulipifera was present and higher condensed and hydrolysable tannins associated with leaf packs lacking L. tulipifera). Macroinvertebrate abundance was consistently lower when R. maximum was present, which was partially explained by litter chemistry [e.g., high concentrations of lignin, condensed tannins, hydrolysable tannins and total phenolics and high carbon to nutrient (N and P) ratios]. 6. The bottom‐up effects of litter species diversity on stream macroinvertebrates and litter breakdown are different, which suggests that structural attributes of macroinvertebrate communities may only partially explain the effects of litter‐mixing on organic matter processing in streams. In addition, stream macroinvertebrates colonising decomposing litter are influenced by resource diversity as well as resource availability. Broad‐scale shifts in riparian tree species composition will alter litter inputs to streams, and our results suggest that changes in the diversity and availability of terrestrial litter may alter stream food webs and organic matter processing.     

Structural features of fragmented woodland communities affect leaf litter decomposition rates

Disruption to the physical structure of plant communities by habitat fragmentation can change microclimates, so leaf litter decomposition rates, being dependent on temperature and moisture, may also be affected. Similarly, smaller-scale structural features of plant communities can modify microclimates, and so may produce distinctive spatial patterns in decomposition rates. We investigated the effects of three types of structural feature having the potential to alter litter layer microclimates: fragmentation-induced modification that diminishes with distance from remnant edges (edge-core); concentric zones of locally modified conditions imposed by individual trees (Belsky–Canham); and highly localised abiotic modification collectively imposed by herbaceous plants (ground cover). We conducted a litter bag experiment in woodland remnants, testing whether the observed spatial variability in litter decomposition was attributable to one or more of these three structural features. The data provided the strongest support for the Belsky–Canham hypothesis, and the least support for the ground cover hypothesis. However, the hypotheses were not mutually exclusive, for each explained a component of the observed variability not explained by either of the other two. Proximity to remnant edge, proximity to trees, canopy light penetration, and ground cover density each explained part of the observed variability between plots. Decomposition rates did not differ with remnant area per se, for the effects of fragmentation were weak, and differed with cardinal direction. In contrast, the effects of individual trees were much stronger, and accounted for most of the between-plot variability. We found that litter decomposition rates in small remnants are only weakly affected by fragmentation, and we consider that the contributions of small remnants to landscape-scale functioning warrant closer attention.     

The influence of humic substances on the aerobic decomposition of submerged leaf litter

Leaf material was incubated in flasks containing streamwater in which the pH and the concentration of isolated fulvic acid were varied independently of one another. Decomposition of the leaf material was slower at pH 4 than at pH 5 or 7, but the concentration of fulvic acid had no effect when the pH was held constant. At pH 5, 20 mg Cl^–1 humic acid also had no effect on decomposition. High concentrations of dissolved fulvic acids may contribute to the slow decomposition of plant litter characteristic of many wetlands through their contribution to hydrogen ion activity, but we could find no evidence for other properties of fulvic acid which inhibit leaf litter decomposition.     

The responses of leaf litter ant communities to wildfires in the Brazilian Amazon: a multi-region assessment

Fire is frequently used as tool for land management in the Amazon, but often escapes into surrounding forests, with potentially severe impacts for forest biodiversity. We investigated the effects of single wildfires on ant communities in four geographically distinct regions of the Brazilian Amazon (Roraima, Pará, Acre and Mato Grosso) where forests had burned between 8 months and 10 years before our sampling. We established 7–12 transects, 500 m each, in burned and unburned forests in each region to investigate the effects of fire on forest structure and leaf litter ant communities, which were sampled using Winkler sacks. Fire effects on forest structure were more drastic in the most recently burned forests in Acre and Mato Grosso, while the impacts of older burns in Roraima and Pará were more subtle. Ant species richness was not different between burnt and unburned areas, but community composition differed between burned and control forests in all regions except Mato Grosso. At the species level, indicator species analysis showed that a limited number of species were significant indicators of unburned control forests in all regions, except Acre. Forests structure variables and leaf litter volume were all important in shaping ant communities, but their relative importance varied between regions. Our results indicate that burned forest have different ant species communities from unburned forests, and those differences are still apparent 10 years after the disturbance, highlighting the importance of effective policies for fire management in Amazon.     

华南4种乡土阔叶树种枯落叶分解能力

采用分解袋法,研究了樟树(Cinnamomum camphora (L.) Presl)、米老排(Mytilaria laosensis Lec.)、火力楠(Michelia macclurei Dandy)和黎蒴(Castanopsis fissa Rehd. et Wils.) 4种华南乡土阔叶树种枯落叶在阔叶混交人工林和湿地松人工林下的分解能力.4个树种枯落叶的分解速率大小依次为:樟树>米老排>火力楠>黎蒴,它们在混交林中的分解速率均大于松林.4个树种枯落叶的分解动态符合Olson指数模型,分解系数(K)在混交林中均大于人工林,且在两种林地中均表现为樟树＞米老排＞火力楠＞黎蒴.不同时间枯落叶养分动态分析结果显示,4个树种枯落叶的N元素分解均出现先富集,后释放的特点,但仅在松林下出现N元素的净释放;K元素释放方式表现为先释放,后富集,仅在阔叶林下出现K元素净释放;Ca、Mg和B元素均表现为分解前期少量富积,之后开始大量释放的变化趋势,它们在阔叶林和松林下均出现净释放.4个树种在P元素释放方式上差异较大,趋势不明显.     

Decomposition of tree leaf litters grown under elevated CO2: Effect of litter quality

Ash (Fraxinus excelsior L.), birch (Betula pubescens Ehrh.), sycamore (Acer pseudoplatanus L.) and Sitka spruce (Picea sitchensis (Bong.) Carr.) leaf litters were monitored for decomposition rates and nutrient release in a laboratory microcosm experiment. Litters were derived from solar domes where plants had been exposed to two different CO₂ regimes: ambient (350 L L^-1 CO₂) and enriched (600 L L^-1 CO₂).Elevated CO₂ significantly affected some of the major litter quality parameters, with lower N, higher lignin concentrations and higher ratios of C/N and lignin/N for litters derived from enriched CO₂. Respiration rates of the deciduous species were significantly decreased for litters grown under elevated CO₂, and reductions in mass loss at the end of the experiment were generally observed in litters derived from the 600 ppm CO₂ treatment. Nutrient mineralization, dissolved organic carbon, and pH in microcosm leachates did not differ significantly between the two CO₂ treatments for any of the species studied. Litter quality parameters were examined for correlations with cumulative respiration and decomposition rates: N concentration, C/N and lignin/N ratios showed the highest correlations, with differences between litter types. The results indicate that higher C storage will occur in soil as a consequence of litter quality changes resulting from higher atmospheric concentrations of CO₂.Abbreviations CHO soluble carbohydrates - DOC dissolved organic carbon - HCel holocellulose - WTREM weight remaining     

Temporal variation overshadows the response of leaf litter microbial communities to simulated global change

Bacteria and fungi drive the decomposition of dead plant biomass (litter), an important step in the terrestrial carbon cycle. Here we investigate the sensitivity of litter microbial communities to simulated global change (drought and nitrogen addition) in a California annual grassland. Using 16S and 28S rDNA amplicon pyrosequencing, we quantify the response of the bacterial and fungal communities to the treatments and compare these results to background, temporal (seasonal and interannual) variability of the communities. We found that the drought and nitrogen treatments both had significant effects on microbial community composition, explaining 2–6% of total compositional variation. However, microbial composition was even more strongly influenced by seasonal and annual variation (explaining 14–39%). The response of microbial composition to drought varied by season, while the effect of the nitrogen addition treatment was constant through time. These compositional responses were similar in magnitude to those seen in microbial enzyme activities and the surrounding plant community, but did not correspond to a consistent effect on leaf litter decomposition rate. Overall, these patterns indicate that, in this ecosystem, temporal variability in the composition of leaf litter microorganisms largely surpasses that expected in a short-term global change experiment. Thus, as for plant communities, future microbial communities will likely be determined by the interplay between rapid, local background variability and slower, global changes.     

The influence of leaf litter on zooplankton in floodplain wetlands: changes resulting from river regulation

1. The loss of input of leaf litter through clearing of riparian vegetation may result in significant changes to aquatic ecosystems. River red gums (Eucalyptus camaldulensis) surrounding floodplain wetlands in the Murray–Darling Basin, Australia, contribute large quantities of leaf litter, but the quality of this resource may change depending on the timing of inundation. 2. We used experimental mesocosms to test the hypotheses that zooplankton would have a greater abundance with an input of leaf litter and that fewer zooplankton would emerge from egg banks in cleared than forested wetlands. The experiment was carried out in summer/autumn and in spring to test a third hypothesis that zooplankton would respond to changes in the timing of wetland inundation as a result of river regulation. 3. In summer/autumn, leaf litter reduced zooplankton abundance by 89% at the beginning of the experiment through its influence on water quality. Only a few taxa (Polyarthra spp., Colurella spp. and the cladoceran Family Moinidae) responded positively to leaf litter when water quality improved later in the experiment, indicating a switch in the role of leaf litter from a non‐trophic to a trophic pathway. 4. In spring, microcrustaceans emerged in smaller numbers from sediment sourced from cleared compared to forested wetlands, reflecting different communities in these two wetland types and/or disturbances to the sediment that interfere with emergence. 5. Although leaf litter appears not to be an important resource for zooplankton in floodplain wetlands, riparian clearing may have lasting effects on future emerging zooplankton communities. Additionally, river regulation may have considerable impacts on the influence of leaf litter on zooplankton, which has implications for the management of floodplain river systems.     

Biodiversity assessment in incomplete inventories: leaf litter ant communities in several types of Bornean rain forest

Biodiversity assessment of tropical taxa is hampered by their tremendous richness, which leads to large numbers of singletons and incomplete inventories in survey studies. Species estimators can be used for assessment of alpha diversity, but calculation of beta diversity is hampered by pseudo-turnover of species in undersampled plots. To assess the impact of unseen species, we investigated different methods, including an unbiased estimator of Shannon beta diversity that was compared to biased calculations. We studied alpha and beta diversity of a diverse ground ant assemblage from the Southeast Asian island of Borneo in different types of tropical forest: diperocarp forest, alluvial forest, limestone forest and heath forests. Forests varied in plant composition, geology, flooding regimes and other environmental parameters. We tested whether forest types differed in species composition and if species turnover was a function of the distance between plots at different spatial scales. As pseudo-turnover may bias beta diversity we hypothesized a large effect of unseen species reducing beta diversity. We sampled 206 ant species (25% singletons) from ten subfamilies and 55 genera. Diversity partitioning among the four forest types revealed that whereas alpha species richness and alpha Shannon diversity were significantly smaller than expected, beta-diversity for both measurements was significantly higher than expected by chance. This result was confirmed when we used the unbiased estimation of Shannon diversity: while alpha diversity was much higher, beta diversity differed only slightly from biased calculations. Beta diversity as measured with the Chao-Sørensen or Morisita-Horn Index correlated with distance between transects and between sample points, indicating a distance decay of similarity between communities. We conclude that habitat heterogeneity has a high influence on ant diversity and species turnover in tropical sites and that unseen species may have only little impact on calculation of Shannon beta diversity when sampling effort has been high.