桂西北喀斯特区原生林与次生林鲜叶和凋落叶化学计量特征

研究喀斯特生态脆弱区植物新鲜叶片与凋落叶的元素化学计量学性状,对该地区森林生态系统的恢复与重建具有重要指导意义。在桂西北喀斯特区分别选取了3个原生林群落与3个次生林群落,研究其建群种植物新鲜叶片和凋落叶的C、N、P元素含量及其生态化学计量特征。结果发现,6个群落建群种新鲜叶片C、N、P含量(其平均含量分别为404.3、22.5、1.75 mg/g)均大于凋落叶(平均含量分别为376.5、19.0、1.35 mg/g),鲜叶C:N、C:P、N:P比值(均值分别为17.8、244.9、13.8)均小于凋落叶(均值分别为19.3、315.3、16.3)。6种植物新鲜叶片N、P含量大于凋落叶,而N:P比小于凋落叶,表明喀斯特区植物对N的再吸收率大于P。3个原生林群落建群种鲜叶与凋落叶的平均C、N含量均大于次生林,而P含量则略小于次生林;原生林鲜叶与凋落叶的C:N比均小于次生林,C:P、N:P则大于次生林,推测次生林相对于原生林有更快的生长速率。原生林鲜叶N:P比为13—15之间,次生林鲜叶N:P比为11—12之间,次生林鲜叶与凋落叶的N:P比均小于原生林,说明原生林凋落物分解相对较慢,原生林能相对多的保留养分以供植物吸收,更能适应喀斯特石生环境。植物鲜叶和凋落叶的C:N与N:P比值均呈极显著正相关,说明叶片养分元素间具有共变的特性;叶片N、P含量呈正相关关系,表明植物N:P比具有相对的稳定性,这是高等陆生植物C-N-P元素计量的普遍规律,体现了植物群落对环境的适应。     

神农架常绿落叶阔叶混交林碳氮磷化学计量比

生态化学计量学是研究生态过程中化学元素平衡的科学, 碳(C)、氮(N)、磷(P)化学计量比是生态系统过程及其功能的重要特征。该研究测定了神农架常绿落叶阔叶混交林植物器官、凋落物及土壤的C、N、P含量, 利用生物量加权法计算其化学计量比, 并分析该生态系统不同组分间及不同器官间化学计量比的差异。研究结果发现: 在不同组分之间, C含量、C:N及C:P表现为植物>凋落物>土壤; N、P含量及N:P表现为凋落物>植物>土壤。在不同植物器官间, C含量的差异较小, 其变异系数相对N、P含量较低且保持稳定; N、P含量为叶片最高且变异系数最低; N:P为树皮最高, 而枝的变异系数最低。常绿与落叶树种的叶片N、P含量差异显著。与不同森林类型的化学计量比相比, 该常绿落叶阔叶混交林植物群落的C:P及N:P较低, 凋落物的C:P及N:P较高, 土壤的C、N、P化学计量比与亚热带常绿阔叶林基本一致, 生态系统的C:N相对较低。利用生物量加权法计算得到的该森林生态系统不同组分的C、N、P化学计量比的大小关系与前人利用枝叶取样算术平均的结果存在较大差异。C、N、P含量及其化学计量比在不同器官的分配及内稳性与器官的生理功能关系密切。     

Characteristics of nitrogen and phosphorus stoichiometry across components of forest ecosystem in Shaanxi Province

Aims Nitrogen (N) and phosphorus (P) stoichiometry between vegetation, litter, and soil were important for understanding biogeochemical cycles in terrestrial ecosystems, but remain poorly understood. Here, our aims were to study characteristics of N and P stoichiometry for the plant, litter, and soil and the interactions between its components across forest ecosystems in Shaanxi Province.Methods A total of 121 sampling sites, covering the most main forest types in Shaanxi, were established across the whole province in 2012. And N and P concentration of vegetation (tree and understory), litter and soil layers were measured for each site.Important findings 1) There were significant differences in the N and P stoichiometry among the forest ecosystem components (p < 0.05). N and P contents were higher in understory and litter layer, and lower in tree and soil. Whereas the N:P was slightly different, highest in litter and lowest in the soil layer, with little differences among remaining components. The contents of N, P and N:P ranged from 0.72 to11.99 mg·g^-1, 0.47 to 1.07 mg·g^-1, and 1.86 to 14.84, respectively. Within top 1 m soil layer N content and N:P decreased with soil depth (p < 0.05), however the P content did not exhibit significant changes. 2) N and P contents, and N:P of each component were higher in broadleaf forest than in coniferous forest, although the difference is not significant. 3) N was positively correlated with P content except for soil and N:P was negatively correlated with P content, but positively correlated with N content within each component. In addition, there was a significant positive correlation for N and P stoichiometric characteristics between litter layer and tree, herb, soil layer. 4) Although spatial pattern of N and P contents, and N:P differed in relation to longitude, latitude, and elevation for forest ecosystem components, a stable distribution was exhibited in general.     

陕西省森林各生态系统组分氮磷化学计量特征

研究森林植被、枯落物和土壤的氮(N)磷(P)化学计量关系对于理解生态系统各组分的相互作用和养分循环具有重要意义。该研究对陕西省不同类型森林生态系统植被、枯落物和土壤的N和P含量及其化学计量关系进行了研究分析。结果表明: 1)森林生态系统各组分的N、P化学计量特征存在显著差异(p < 0.05), N、P含量均以林下灌草层植物和枯落物层较高, 乔木层植物和土壤层较低; N:P值则稍有不同, 以枯落物层最高, 土壤层最低, 其他各层差异不显著; 各组分N、P含量和N:P值分别为0.72-11.99 mg·g^-1、0.47-1.07 mg·g^-1和1.86-14.84。0-1 m土层内N含量、N:P值均随土层加深而降低(p < 0.05), P含量则不随土层发生明显变化。2)各组分N、P含量和N:P值多表现为阔叶林高于针叶林, 但其差异不显著。3)生态系统同一组分内, N、P含量间极显著正相关, N、P含量与N:P值分别呈极显著正相关、负相关关系, 但是土壤层内N、P含量无显著相关关系。各组分间, 枯落物层与乔木层、草本层和土壤层的N、P含量和N:P值也均极显著正相关, 而枯落物层与灌木层植物无显著相关关系。4)生态系统各组分N、P含量和N:P值随空间变化表现不尽一致, 总体上呈稳态。该文通过对N、P化学计量特征的研究, 揭示了森林生态系统植被、枯落物和土壤组分间所存在的养分循环联系, 这些联系中也表现出分异特征, 而分异可能由各自所执行的不同生态功能所致。     

不同林龄油松(Pinus tabulaeformis)人工林植物、凋落物与土壤C、N、P化学计量特征

在陕西省北部延安市境内子午岭林区,采用时空互代的方法选取9、23、33、47年生油松(Pinus tabuliformis)人工林为研究对象,比较油松不同器官(叶、枝、干、根)、凋落物及土壤C、N、P含量及其比值的差异,探讨它们随林龄的变化及其相互间的关系,以期为油松人工林的生产、改善和林木生长环境的调节提供参考。结果表明:除根中C含量在林龄间差异不显著外,其它器官C、N、P含量及其比值在林龄间均差异显著且随林龄增加变化趋势不尽相同。9、23、33、47年生油松林C、N、P含量及N∶P比值均在叶中最高;C∶N比值均在干中最高,根中次之;C∶P比值均在干中最高,其它器官大小次序不一。除33年生油松林叶中N∶P比值大于14外,其它各器官各林龄N∶P比值均小于14,且N∶P比值随林龄先增加后减少,故可判断油松在该区域受N限制较为严重,且随林龄的增加受N限制的情况有所缓解。不同林龄土壤和凋落物C、N、P含量及其比值差异显著,且后者均大于前者。土壤与凋落物C、P含量及C∶N、C∶P、N∶P比值随林龄增加变化趋势完全一致,表明土壤与凋落物之间有着密切的关系。叶片与凋落物N、P含量及C∶N、C∶P、N∶P比值之间显著相关,表明凋落物的养分承自植物叶片,二者之间关系紧密;植物和土壤的C、N、P含量之间均不存在显著相关性,说明土壤C、N、P供应量对乔木叶片C、N、P含量影响不大。     

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.     

陕西省森林群落乔灌草叶片和凋落物C、N、P生态化学计量特征

研究陕西省121个森林群落中乔叶、灌叶、草叶和凋落物4个层次的C、N、P化学计量学特征及其变异性,分析了各层次间及其与地理因子间的关系.结果表明： 乔叶C、N含量最高,草叶P含量最高,凋落物C、N、P含量均最低.乔叶、灌叶、草叶和凋落物的C∶N∶P分别为439.4∶14.2∶1、599.2∶13.5∶1、416.5∶13.3∶1、504.8∶15.5∶1,乔叶、灌叶、草叶N∶P值差异不显著,但均与凋落物N∶P值差异显著,表明不同生活型活体植物叶片的N∶P值具有一定的稳定性.与N含量相比,C和P含量在各层次间表现出更好的相关性;C、N、P含量及其比值在草叶与乔叶之间较在灌叶与乔叶之间表现出更好的相关性;凋落物与乔叶和草叶之间C、N、P含量均表现出显著正相关,与灌叶只在P含量之间存在显著正相关.灌叶N、P含量随纬度增加而增加,草叶N、P含量随经度增加而降低,海拔对叶片和凋落物C、N、P含量及其比值的影响较小,只有乔叶N∶P与海拔之间存在显著负相关.各地理因子对草叶的影响大小表现为经度>纬度>海拔,对乔叶、灌叶和凋落物的影响大小表现为纬度>经度>海拔.     

Carbon : nitrogen stoichiometry in forest ecosystems during stand development

Aim Carbon (C) and nitrogen (N) stoichiometry is a critical indicator of biogeochemical coupling in terrestrial ecosystems. However, our current understanding of C : N stoichiometry is mainly derived from observations across space, and little is known about its dynamics through time. Location Global secondary forests. Methods We examined temporal variations in C : N ratios and scaling relationships between N and C for various ecosystem components (i.e. plant tissue, litter, forest floor and mineral soil) using data extracted from 39 chronosequences in forest ecosystems around the world. Results The C : N ratio in plant tissue, litter, forest floor and mineral soil exhibited large variation across various sequences, with an average of 145.8 ± 9.4 (mean ± SE), 49.9 ± 3.0, 38.2 ± 3.1 and 18.5 ± 0.9, respectively. In most sequences, the plant tissue C : N ratio increased significantly with stand age, while the C : N ratio in litter, forest floor and mineral soil remained relatively constant over the age sequence. N and C scaled isometrically (i.e. the slope of the relationship between log‐transformed N and C is not significantly different from 1.0) in litter, forest floor and mineral soil both within and across sequences, but not in plant tissue either within or across sequences. The C : N ratio was larger in coniferous forests than in broadleaf forests and in temperate forests than in tropical forests. In contrast, the N–C scaling slope did not reveal significant differences either between coniferous and broadleaf forests or between temperate and tropical forests. Main conclusions These results suggest that C and N become decoupled in plants but remain coupled in other ecosystem components during stand development.     

N limitation increases along a temperate forest succession: evidences from leaf stoichiometry and nutrient resorption

温带森林演替加剧了氮限制：来自叶片化学计量和养分重吸收的证据 森林生产力和碳汇功能在很大程度上取决于土壤氮和磷的有效性。然而,迄今为止,养分限制随森林演替的时间变化仍存在争议。叶片化学计量和养分重吸收是预测植物生长养分限制的重要指标。基于此,本研究测定了温带森林4个演替阶段所有木本植物叶片和凋落叶中氮和磷的含量,并分析了演替过程中非生物因子和生物因子如何影响叶片化学计量和养分重吸收。研究结果表明,在个体尺度上,叶片氮磷含量在演替末期显著增加,而叶片氮磷比无显著变化;氮的重吸收效率随演替显著增加,然而磷的重吸收效率先增加后减少;氮重吸收效率与磷重吸收效率的比值仅在演替末期显著增加。此外,植物氮素循环对土壤养分的响应比磷素循环更弱。在群落尺度上,叶片氮磷含量随森林演替呈现先降低后升高的趋势,主要受香农-维纳多样性指数和物种丰富度的影响;叶片氮磷比随演替而显著变化,主要由胸径的群落加权平均值决定;氮的重吸收效率增加,主要受物种丰富度和胸径的影响,而磷的重吸收效率相对稳定。因此,氮重吸收效率与磷重吸收效率的比值显著增加,表明随着温带森林演替,氮限制加剧。这些结果可能反映了较高生物多样性群落中物种间对有限资源的激烈竞争,强调了生物因子在驱动森林生态系统养分循环中的重要性,为中国温带和北方森林可持续经营的施肥管理提供了参考。     

Leaf litter carbon,nitrogen, and phosphorus stoichiometric patterns as related to climatic factors and leaf habits across Chinese broad-leaved tree species

Our understanding of how climate and leaf habit (evergreen vs. deciduous) drive leaf litter carbon (C), nitrogen (N), and phosphorus (P) stoichiometric patterns is largely limited but is particularly important for broad-leaved forests, since the forest is sensitive to climate change. Here, we investigated leaf litter C, N, and P stoichiometric ratios of broad-leaved tree species in relation to climate and leaf habit using previous publications and our additional samplings across China. We found that mean leaf litter C:N across Chinese broad-leaved tree species was within the range of the global flora, whereas C:P was lower and N:P higher. Evergreen species displayed higher leaf litter C:N, C:P, and N:P than their deciduous counterparts. Both leaf litter C:P and N:P for all species pooled were negatively correlated with latitude, driven by mean annual precipitation (MAP) and mean annual temperature, respectively, while leaf litter C:N displayed no clear latitudinal trend. The direction and magnitude of leaf litter C, N, and P stoichiometric ratios in response to climate diverged between leaf habits. For example, evergreen leaf litter C:N was negatively correlated with MAP, while deciduous counterparts did not respond significantly to MAP. We conclude that leaf litter C, N, and P stoichiometric ratios shifted along the climatic gradient, and the strength of such shifts differed between leaf habits. Therefore, leaf litter stoichiometric patterns across leaf habits suggest that any climate change-driven shift in species distribution may potentially alter the ecosystem’s nutrient cycling processes of evergreen- and deciduous-dominated broad-leaved forests differentially.     

Variations in leaf litter decomposition across contrasting forest stands and controlling factors at local scale

Aims Litter decomposition is a critical pathway linking the above- and belowground processes. However, factors underlying the local spatial variations in forest litter decomposition are still not fully addressed. We investigated leaf litter decomposition across contrasting forest stands in central China, with objective to determine the spatial variations and controlling factors in forest floor leaf litter decomposition in relation to changes in forest stands in a temperate forest ecosystem.Methods Leaf litter decomposition was studied by using litterbag method across several typical forest stand types in Baotianman Nature Reserve, central China, including pure stands of Quercus aliena var. acuteserrata, Q. glandulifera var. brevipetiolata and Q. variabilis, respectively, and mixed pine/oak stands dominated by Pinus armandii and Q. aliena var. acuteserrata, as well as stands of pure Q. aliena var. acuteserrata trees ranging in stand age from ~40 to>160 years. Measurements were made on litter mass remaining and changes in litter chemistry during decomposition over a 2-year period, along with data collections on selective biotic and environmental factors. A reciprocal transplant experiment involving Q. aliena var. acuteserrata and Q. variabilis was concurrently carried out to test the occurrence of 'home-field advantage (HFA)' in local forests when only considering contrasting oak tree species. Correlation analyses and path analyses were performed to identify the dominant drivers and their relative contributions to variations in leaf litter decomposition.Important findings Significant variations were found in the rate of leaf litter decomposition among stands of different tree species but not among stand age classes. The values of decay constant, k, varied from 0.62 in Q. aliena var. acuteserrata stands to 0.56 in Q. variabilis stands. The reciprocal litter transplant experiment showed that the rate of leaf litter decomposition was on average 5% slower in home-fields than on reciprocal sites. Path analysis identified litter acid-unhydrolyzable residue (AUR) to N ratio, soil microbial biomass carbon (MBC), soil pH and soil organic carbon (SOC) as most prominent factors controlling the rate of leaf litter decomposition, collectively accounting for 57.8% of the variations; AUR/N had the greatest negative effect on k value, followed by weaker positive effects of SOC and MBC. Our findings suggest that tree species plays a primary role in affecting forest floor leaf litter decomposition by determining the litter quality, with site environment being a secondary factor contributing to the local variations in leaf litter decomposition in this temperate forest ecosystem.     

大兴安岭典型森林沼泽植物叶片和细根碳氮磷化学计量特征

探究大兴安岭典型森林沼泽不同植物叶片和细根生态化学计量特征,能够为进一步认识高纬度气候敏感生态系统养分利用策略和物质循环过程提供依据。对大兴安岭地区兴安落叶松-苔草、兴安落叶松-笃斯越桔-藓类和兴安落叶松-杜香-泥炭藓3种典型森林沼泽19种优势和亚优势维管植物叶片和细根碳氮磷计量特征(C∶N∶P)进行比较,分析不同森林沼泽类型、植物生长型和菌根类型叶片和细根C∶N∶P差异,通过标准化主轴回归分析叶片与细根C∶N∶P的关系。结果表明: 叶片C∶N∶P在种间水平具有最大的变异(42.5%～84.6%),且叶片和细根种间变异大小均为N∶P>C∶N>C∶P。土壤养分和水分含量较高的兴安落叶松-苔草沼泽叶片与细根C∶N和C∶P值较低,且3种森林沼泽植物叶片和细根N∶P均小于10,受N限制。草本植物叶片C∶P和细根C∶N、C∶P显著低于木本植物。外生菌根和杜鹃花类菌根植物叶片和细根C∶N和C∶P高于丛枝菌根和无菌根植物,且杜鹃花类菌根植物叶片和细根C∶P显著高于外生菌根植物。不同森林沼泽、生长型、菌根类型植物叶片和细根C∶N和C∶P差异明显,而N∶P相对稳定。森林沼泽植物叶片与细根C∶N、C∶P和N∶P呈线性正相关,植物地上与地下部分在生态化学计量特征上存在协同。     

Detritivore stoichiometric diversity alters litter processing efficiency in a freshwater ecosystem

Many studies have estimated relationships between biodiversity and ecosystem functioning, and observed generally positive effects. Because detritus is a major food resource in stream ecosystems, decomposition of leaf litter is an important ecosystem process and many studies report the full range of positive, negative and no effects of diversity on decomposition. However, the mechanisms underlying decomposition processes in fresh water remain poorly understood. Organism body stoichiometry relates to consumption rates and tendencies, and decomposition processes of litter may therefore be affected by diversity in detritivore body stoichiometry. We predicted that the stoichiometric diversity of detritivores (differences in C: nutrient ratios among species) would increase the litter processing efficiency (litter mass loss per total capita metabolic capacity) in fresh water through complementation regarding different nutrient requirements. To test this prediction, we conducted a microcosm experiment wherein we manipulated the stoichiometric diversity of detritivores and quantified mass loss of leaf litter mixtures. We compared litter processing efficiency among litter species in each microcosm with single species detritivores, and observed detritivores with nutrient‐rich bodies tended to prefer litter with lower C: nutrient ratios over litter with higher C: nutrient ratios. Furthermore, litter processing efficiencies were significantly higher in the microcosms containing species of detritivores with both nutrient‐rich and ‐poor bodies than microcosms containing species of detritivores including only nutrient‐rich or ‐poor bodies. This might mean a higher stoichiometric diversity of detritivores increased litter processing efficiency. Our results suggest that ecological stoichiometry may improve understanding of links between biodiversity and ecosystem function in freshwater ecosystems.     

High variation in foliage and leaf litter chemistry among 45 tree species of a neotropical rainforest community

Distinct ecosystem level carbon : nitrogen : phosphorus (C : N : P) stoichiometries in forest foliage have been suggested to reflect ecosystem-scale selection for physiological strategies in plant nutrient use. Here, this hypothesis was explored in a nutrient-poor lowland rainforest in French Guiana. Variation in C, N and P concentrations was evaluated in leaf litter and foliage from neighbour trees of 45 different species, and the litter concentrations of major C fractions were also measured. Litter C ranged from 45.3 to 52.4%, litter N varied threefold (0.68-2.01%), and litter P varied seven-fold (0.009-0.062%) among species. Compared with foliage, mean litter N and P concentrations decreased by 30% and 65%, respectively. Accordingly, the range in mass-based N : P shifted from 14 to 55 in foliage to 26 to 105 in litter. Resorption proficiencies indicated maximum P withdrawal in most species, but with a substantial increase in variation in litter P compared with foliage. These data suggest that constrained ecosystem-level C : N : P ratios do not preclude the evolution of highly diversified strategies of nutrient use and conservation among tropical rainforest tree species. The resulting large variation in litter quality will influence stoichiometric constraints within the decomposer food web, with potentially far-ranging consequences on nutrient dynamics and plant-soil feedbacks.     

Tree leaf litter composition and nonnative earthworms influence plant invasion in experimental forest floor mesocosms

Dominant tree species influence community and ecosystem components through the quantity and quality of their litter. Effects of litter may be modified by activity of ecosystem engineers such as earthworms. We examined the interacting effects of forest litter type and earthworm presence on invasibility of plants into forest floor environments using a greenhouse mesocosm experiment. We crossed five litter treatments mimicking historic and predicted changes in dominant tree composition with a treatment of either the absence or presence of nonnative earthworms. We measured mass loss of each litter type and growth of a model nonnative plant species (Festuca arundinacea, fescue) sown into each mesocosm. Mass loss was greater for litter of tree species characterized by lower C:N ratios. Earthworms enhanced litter mass loss, but only for species with lower C:N, leading to a significant litter × earthworm interaction. Fescue biomass was significantly greater in treatments with litter of low C:N and greater mass loss, suggesting that rapid decomposition of forest litter may be more favorable to understory plant invasions. Earthworms were expected to enhance invasion by increasing mass loss and removing the physical barrier of litter. However, earthworms typically reduced invasion success but not under invasive tree litter where the presence of earthworms facilitated invasion success compared to other litter treatments where earthworms were present. We conclude that past and predicted future shifts in dominant tree species may influence forest understory invasibility. The presence of nonnative earthworms may either suppress of facilitate invasibility depending on the species of dominant overstory tree species and the litter layers they produce.     

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。     

Non‐symbiotic nitrogen fixation during leaf litter decomposition in an old‐growth temperate rain forest of Chiloé Island,southern Chile: Effects of single versus mixed species litter

Heterotrophic nitrogen fixation is a key ecosystem process in unpolluted, temperate old‐growth forests of southern South America as a source of new nitrogen to ecosystems. Decomposing leaf litter is an energy‐rich substrate that favours the occurrence of this energy demanding process. Following the niche ‘complementarity hypothesis’, we expected that decomposing leaf litter of a single tree species would support lower rates of non‐symbiotic N fixation than mixed species litter taken from the forest floor. To test this hypothesis we measured acetylene reduction activity in the decomposing monospecific litter of three evergreen tree species (litter C/N ratios, 50–79) in an old‐growth rain forest of Chiloé Island, southern Chile. Results showed a significant effect of species and month (anova , Tukey's test, P < 0.05) on decomposition and acetylene reduction rates (ARR), and a species effect on C/N ratios and initial % N of decomposing leaf litter. The lowest litter quality was that of Nothofagus nitida (C/N ratio = 78.7, lignin % = 59.27 ± 4.09), which resulted in higher rates of acetylene reduction activity (mean = 34.09 ± SE = 10.34 nmol h^?1 g^?1) and a higher decomposition rate (k = 0.47) than Podocarpus nubigena (C/N = 54.4, lignin % = 40.31 ± 6.86, Mean ARR = 4.11 ± 0.71 nmol h^?1 g^?1, k = 0.29), and Drimys winteri (C/N = 50.6, lignin % = 45.49 ± 6.28, ARR = 10.2 ± 4.01 nmol h^?1 g^?1, k = 0.29), and mixed species litter (C/N = 60.7, ARR = 8.89 ± 2.13 nmol h^?1g^?1). We interpret these results as follows: in N‐poor litter and high lignin content of leaves (e.g. N. nitida) free‐living N fixers would be at competitive advantage over non‐fixers, thereby becoming more active. Lower ARR in mixed litter can be a consequence of a lower litter C/N ratio compared with single species litter. We also found a strong coupling between in situ acetylene reduction and net N mineralization in surface soils, suggesting that as soon N is fixed by diazotroph bacteria it may be immediately incorporated into mineral soil by N mineralizers, thus reducing N immobilization.     

Stoichiometric imbalances between detritus and detritivores are related to shifts in ecosystem functioning

How are resource consumption and growth rates of litter‐consuming detritivores affected by imbalances between consumer and litter C:N:P ratios? To address this question, we offered leaf litter as food to three aquatic detritivore species, which represent a gradient of increasing body N:P ratios: a crustacean, a caddisfly and a stonefly. The detritivores were placed in microcosms and submerged in a natural stream. Four contrasting leaf species were offered, both singly and in two‐species mixtures, to obtain different levels of stoichiometric imbalance between the resources and their consumers. The results suggest that detritivore growth was constrained by N rather than C or P, even though 1) the N:P ratios of the consumers’ body tissue was relatively low and 2) microbial leaf conditioning during the experiment reduced the N:P imbalance between detritivores and leaf litter. This surprisingly consistent N limitation may be a consequence of cumulative N‐demand arising from the production of N‐rich chitin in the exoskeletons of all three consumer species, which is lost during regular moults, in addition to N‐demand for silk production by the caddisfly. These N requirements are not commonly quantified in stoichiometric analyses of arthropod consumers. There was no evidence for compensatory feeding, but when offered mixed‐species litter varying in C:N:P ratios, detritivores consumed more of the litter species showing the highest N:P and lowest C:N ratio, accelerating the mass loss of the preferred leaf species in the litter mixture. These results show that imbalances in consumer–resource stoichiometry can have contrasting effects on coupled processes, highlighting a challenge in developing a mechanistic understanding of the role of stoichiometry in regulating ecosystem processes such as leaf litter decomposition.     

青藏高原东缘主要针叶树种叶片碳氮磷化学计量分布格局及其驱动因素

理解植物叶片化学计量特征及其驱动因素对认识植物种群分布规律及预测植物对环境变化响应具有重要意义。该研究采集了青藏高原东缘针叶林84个样点共29种主要针叶树种叶片, 探讨该区域常绿针叶树种叶片碳(C)、氮(N)、磷(P)化学计量特征和分布格局及其驱动因素。结果表明: (1)在科和属水平上, 不同针叶树种叶片C、N含量和C:N差异显著; 叶片N:P < 14, 表明该区域针叶树种主要受N限制。(2)叶片N、P含量在环境梯度上表现出一致的分布规律: 均呈现出随纬度和海拔增加而显著降低, 随年平均气温(MAT)和年降水量(MAP)增加而显著增加的趋势; 而叶片C含量与纬度、海拔、MAT和MAP均未表现出显著相关性。(3)叶片C:N、C:P呈现出与N、P含量变化相反的分布格局: 均随纬度和海拔增加而显著增加, 随MAT和MAP增加而显著降低; 而叶片N:P与海拔、MAT和MAP均无显著相关性。(4)进一步分析表明, 叶片C、N、P含量及其化学计量比的主要驱动因素不尽相同。具体而言: 土壤特性是叶片C含量和N:P变异的主要驱动因子, 而叶片N、P含量和C:N、C:P的变异主要由气候因素决定。总之, 该区域针叶树种叶片化学计量沿环境梯度的变异规律有力地支持了温度生物地球化学假说, 在一定程度上丰富了对环境变化下植物叶片化学计量分布格局及其驱动机制的认识。     

Impacts of nitrogen addition on foliar nitrogen and phosphorus stoichiometry in a subtropical evergreen broad-leaved forest in Mount Wuyi

Aims Our purpose was to explore the effects of nitrogen addition on foliar nitrogen (N), phosphorus (P) and N:P stoichiometry and to assess their differences among different species and functional groups.
Methods N addition experiment has been conducted in a subtropical evergreen broad-leaved forest in Mount Wuyi, Fujian Province since 2011. Foliar concentrations of nitrogen and phosphorus were measured and foliar stoichiometry was estimated in tree, shrub, herb, fern and moss species following the N addition treatments from 2013 to 2015.
Important findings Generally, foliar N increased for almost all species and herbaceous plants are much more sensitive than trees and shrubs under N addition. Foliar N of Castanopsis carlesii, Amomum villosum, Woodwardia japonica increased significantly under N addition. Foliar P for most species was sensitive to the N addition. Foliar P of herbaceous plants increased significantly but foliar P of Leucobryum chlorophyllosum decreased significantly. The results showed the subtropical evergreen forest in Mount Wuyi was mainly limited by P and mean foliar N:P ratios enhanced from 18.67 to 19.72 under N addition, indicating that the strength of P limitation was enhanced by N addition. N:P ratios of the dominant arboreal species in the communities tended to be stable, while N:P ratios of herbaceous plants and shrubs increased. The changes in N:P ratios were mainly determined by P dynamics instead of N dynamics under N addition, and our results confirmed that increasing N availability can affect P cycling.