森林凋落物分解研究进展

森林凋落物是指森林生态系统内由生物组分产生,然后归还到林地表面的所有有机物质的总称。森林凋落物在促进森林生态系统正常的物质循环和养分平衡,维持生态系统功能中具有重要作用,其分解受多因素影响,且各因素之间相互交错。不同情况下,各因子的重要性可能不同。温度和湿度被认为是影响凋落物分解主要的气候因子。凋落物随着温度升高分解速率加快,增加土壤湿度对凋落物分解有积极作用。凋落物的化学性质中,C、N比和木质素含量被认为是最重要的指标。凋落物分解前期的分解速率受到养分含量、水溶性碳化合物和结构碳化合物含量的强烈影响,而后期则更多地受到木质素及纤维素/木质素比值的支配。土壤动物可以粉碎凋落物,土壤微生物也是促进凋落物分解的重要因素,人为活动也影响凋落物分解。N沉降、全球变暖和臭氧层破坏等全球变化对森林凋落物分解的影响已逐渐成为研究热点。未来凋落物分解的研究方向是统一研究方法,开展长期定位监测,加强对分解过程中有机碳含量和释放量的研究,以及N沉降对凋落物分解作用机理的研究。     

Assessing the carbon balance of circumpolar Arctic tundra using remote sensing and process modeling.

This paper reviews the current status of using remote sensing and process-based modeling approaches to assess the contemporary and future circumpolar carbon balance of Arctic tundra, including the exchange of both carbon dioxide and methane with the atmosphere. Analyses based on remote sensing approaches that use a 20-year data record of satellite data indicate that tundra is greening in the Arctic, suggesting an increase in photosynthetic activity and net primary production. Modeling studies generally simulate a small net carbon sink for the distribution of Arctic tundra, a result that is within the uncertainty range of field-based estimates of net carbon exchange. Applications of process-based approaches for scenarios of future climate change generally indicate net carbon sequestration in Arctic tundra as enhanced vegetation production exceeds simulated increases in decomposition. However, methane emissions are likely to increase dramatically, in response to rising soil temperatures, over the next century. Key uncertainties in the response of Arctic ecosystems to climate change include uncertainties in future fire regimes and uncertainties relating to changes in the soil environment. These include the response of soil decomposition and respiration to warming and deepening of the soil active layer, uncertainties in precipitation and potential soil drying, and distribution of wetlands. While there are numerous uncertainties in the projections of process-based models, they generally indicate that Arctic tundra will be a small sink for carbon over the next century and that methane emissions will increase considerably, which implies that exchange of greenhouse gases between the atmosphere and Arctic tundra ecosystems is likely to contribute to climate warming.     

Long-term forest-floor litter dynamics in Canada's boreal forest: Comparison of two model formulations

Litter decomposition is a key component in ecosystem material cycling that determines (i) forest soil carbon (C) and nutrient content, (ii) release of carbon dioxide to the atmosphere, and (iii) generation and mass transfer of dissolved organic carbon from terrestrial to aquatic ecosystems. In this study, we provide simulations of long term forest-floor litter dynamics generated with both (i) an existing forest nutrient cycling and biomass growth model (ForNBM) with a single-pool formulation of forest-floor litter decomposition (Zhu et al., 2003. Ecol. Model. 169, 347-360), and (ii) a revised version of the model produced by substituting the single-pool formulation with a three-pool version of the formulation tested against data from litterbag experiments (FLDM; Zhang et al., 2010. Ecol. Model. 221, 1944-1953). This is done to determine the importance of subdividing the litter mass into categories of rates of decay (i.e., fast, slow, and very slow) on model accuracy. Forest-forest litter dynamics simulated with the two models are subsequently compared against field measurements collected at several northern jack pine (Pinus banksiana) stands along a southwest-northeast oriented transect (climate gradient) associated with the Boreal Forest Transect Case Study in northwest Canada. Initial comparison shows that the single-pool formulation underpredicts residual litter mass when forests are <65 years old, largely due to the improper treatment of the very slow decomposing litter component. This underprediction is resolved when the three-pool formulation is used. From a ecosystems-response point of view, the revised ForNBM (with the three-pool formulation) demonstrates that (i) forest-floor litter initially increases with forest growth and reaches a plateau once the forest matures; (ii) the forest floor stores more litter and C at the southern and warmer sites than at the northern sites; and (iii) in a similar climate regime, the forest floor stores more litter and C at productive than at nutrient-poor sites.     

Nutrient regulation of organic matter decomposition in a tropical rain forest

Terrestrial biosphere-atmosphere CO2 exchange is dominated by tropical forests, so understanding how nutrient availability affects carbon (C) decomposition in these ecosystems is central to predicting the global C cycle's response to environmental change. In tropical rain forests, phosphorus (P) limitation of primary production and decomposition is believed to be widespread, but direct evidence is rare. We assessed the effects of nitrogen (N) and P fertilization on litter-layer organic matter decomposition in two neighboring tropical rain forests in southwest Costa Rica that are similar in most ways, but that differ in soil P availability. The sites contain 100-200 tree species per hectare and between species foliar nutrient content is variable. To control for this heterogeneity, we decomposed leaves collected from a widespread neotropical species, Brosimum utile. Mass loss during decomposition was rapid in both forests, with B. utile leaves losing >80% of their initial mass in <300 days. High organic matter solubility throughout decomposition combined with high rainfall support a model of litter-layer decomposition in these rain forests in which rapid mass loss in the litter layer is dominated by leaching of dissolved organic matter (DOM) rather than direct CO2 mineralization. While P fertilization did not significantly affect mass loss in the litter layer, it did stimulate P immobilization in decomposing material, leading to increased P content and a lower C:P ratio in soluble DOM. In turn, increased P content of leached DOM stimulated significant increases in microbial mineralization of DOM in P-fertilized soil. These results show that, while nutrients may not affect mass loss during decomposition in nutrient-poor, wet ecosystems, they may ultimately regulate CO2 losses (and hence C storage) by limiting microbial mineralization of DOM leached from the litter layer to soil.     

A process-based model of nitrogen cycling in forest plantations: Part I. Structure,calibration and analysis of the decomposition model

We present a new decomposition model of C and N cycling in forest ecosystems that simulates N mineralisation from decomposing tree litter. It incorporates a mechanistic representation of the role of soil organisms in the N mineralisation-immobilisation turnover process during decomposition. We first calibrate the model using data from decomposition of ¹⁴C-labelled cellulose and lignin and ¹⁴C-labelled legume material and then calibrate and test it using mass loss and N loss data from decomposing Eucalyptus globulus residues. The model has been linked to the plant production submodel of the G’DAY ecosystem model, which previously used the CENTURY decomposition submodel for simulating C and N cycling. The key differences between this new decomposition model and the previous one, based on the CENTURY model, are: (1) growth of microbial biomass is the process that drives N mineralisation-immobilisation, and microbial succession is simulated; (2) decomposition of litter can be N-limited, depending on soil inorganic N availability relative to N requirements for microbial growth; (3) ‘quality’ of leaf and fine root litter is expressed in terms of biochemically measurable fractions; (4) the N:C ratio of microbial biomass active in decomposing litter is a function of litter quality and N availability; and (5) the N:C ratios of soil organic matter (SOM) pools are not prescribed but are instead simulated output variables defined by litter characteristics and soil inorganic N availability. With these modifications the model is able to provide reasonable estimates of both mass loss and N loss by decomposing E. globulus leaf and branch harvest residues in litterbag experiments. A sensitivity analysis of the decomposition model to selected parameters indicates that parameters regulating the stabilisation of organic C and N, as well as those describing incorporation of soil inorganic N in Young-SOM (biochemical immobilisation of N) are particularly critical for long-term applications of the model. A parameter identifiability analysis demonstrates that simulated short-term C and N loss from decomposing litter is highly sensitive to three model parameters that are identifiable from the E. globulus litterbag data.     

气候变化与土壤有机质的关系(英文)

在最近的地质历史中,气候变化的速率是史无前例的,它对地球生物圈产生了巨大的影响。土壤有机质中的碳是地球碳库的重要组成部分,它参与全球碳循环。土壤有机质分解而产生的CO2和CH4是重要的温室气体。土壤有机质对气候模型的反应较敏感;其总量取决于生物量生产与分解的平衡状态,以及土壤储存有机质的能力。就全球规模来说,土壤有机质沿着降水增加和温度下降的梯度而增加。温度是支配凋落物分解速率的重要环境因素,它甚至能改变凋落物分解的动力学。     

森林土壤有机层生化特性及其对气候变化的响应研究进展

森林土壤有机层是指累积在土壤表面未分解到完全分解的有机残余物质,在全球碳循环中具有十分重要的作用和地位.目前有关森林土壤有机层的生态研究主要集中于土壤有机层的凋落物储量、水土保持功能、生物多样性保育功能及其生化特性等,而有关其对气候变化响应的研究报道还相当少见,且已有的研究主要关于土壤有机层的碳源/汇动态等,有关森林土壤有机层生化特性对气候变化响应的研究还相对较少,这与其在全球气候变化中的作用和地位是极不相称的.过去10a中,有关土壤有机层生化特性对气候变化响应的研究主要包括土壤有机层的微生物数量、微生物生物量、呼吸作用、有机物质分解动态(凋落物分解)、酶活性等对环境变化的响应等方面.进一步的控制实验研究被认为是相当重要的.参51     

鼎湖山马尾松林凋落物分解对凋落物输入变化的响应

为了了解我国南方森林常见的人为干扰（凋落物收取）活动对生态系统养分循环的影响,研究了鼎湖山马尾松林3种主要树种凋落物分解及其养分释放对凋落物输入量变化的响应。这3种树种分别为马尾松（Pinus massoniana）、荷木（Schimasuperba）和锥栗（Castanopsis chinensis）。凋落物输入量变化分别为凋落物去除（L-）、加倍（L＋）和对照（L）3种处理,每种处理25个重复。经过18个月的处理试验,凋落物分解速率及其养分释放随树种、分解阶段和凋落物处理不同而异。荷木、马尾松和锥栗分解物平均残留率分别为0.46±0.01、0.42±0.01、0.40±0.02,其中,荷木与锥栗、马尾松差异性显著。不同处理间的凋落物分解速率差异显著,加倍、对照和去除处理样地凋落物的平均残留率分别为0.51±0.08、0.53±0.09和0.55±0.08。凋落物加倍处理促进了凋落物分解过程中C的释放,而去除凋落物处理则抑制了N、P的释放。以上结果表明,凋落物收取活动不仅直接带走凋落物中的大量养分,而且抑制了凋落物分解及其养分释放。     

Indirect effects of soil invertebrates on litter decomposition: Elaboration via analysis of a tundra model

Indirect effects of soil invertebrates on litter decomposition in arctic coastal tundra were examined by means of a cross control option in a linear carbon flow model. Systems analysis and simulated field experiments generated predictions about soil systems containing invertebrate effects as modeled. Fauna were found to be several times more important per gram than microflora in causing higher cycling efficiency and more even resource distribution within the model. Detritus enrichment decreased evennes of resource distribution without affecting cycling efficiency. Hypotheses were generated which, if supported by field data, would help confirm the rationale used in model construction.     

Effects of soil nematodes communities on litter decomposition in urban forest of Dalian北大核心CSCD

In order to further expound the effect of soil nematodes on litter decomposition in urban forest, this study investigated the responses of soil nematodes communities to litter decomposition by litterbags technology at different mesh sizes in Dalian National Forest Park. Soil nematodes community composition, decomposition rate of litter and nutrient release were also analyzed. It found a total of 4 418 nematodes from 39 genera. Higher relative density of soil nematodes was found in the bags with 0.1 mm mesh size (1.55 individuals and 7.34 orders per g of dry litter) compared to the bags with 0.02 mm mesh size (0.21 individuals and 0.49 orders per g of dry litter). There were very few soil nematodes in the 0.02 mm litter bags. Thus we regarded that soil nematodes only affected the 0.1 mm litter bags. The mass loss rates as well as C and P release rates was higher in litter bags with 0.1 mm mesh size than in those with 0.02 mm mesh size, indicating a significant influence of soil nematodes on mass loss and nutrient release. The contribution of soil nematodes to the litter mass loss was about 24%. The effect of soil nematodes on the nutrient release rates differed among elements. The soil nematodes had a bigger influence on the release rate of N than that of other elements, whereas the influence on K release rate was the lowest. The results showed that soil nematodes communities has a significant effect on the litter decomposition and nutrient release of the forest litter in Dalian city, especially promoting the mass loss rates and N release rates.     

模拟氮沉降对温带森林凋落物分解的影响

森林凋落物的分解是生态系统养分循环的重要过程,以北京西山地带性植被栎树林（辽东栎：Quercus liaotungensis）为对象,主要研究温带森林植物凋落物分解对模拟氮沉降的响应,为更好地了解氮沉降对温带森林地区凋落物的分解过程提供参考.通过模拟氮沉降,研究不同形态氮（硝态氮、铵态氮和混合态氮）和不同水平氮沉降（对照0 kg·hm^-2·a^-1、低氮处理50 kg·hm^-2·a^-1 和高氮处理150 kg·hm^-2·a^-1）对凋落物分解的影响,在2 年的时间内调查分析了凋落物分解过程中质量损失动态和碳（C）、N 含量及w（C）/w（N）比值的变化.研究结果表明,氮沉降均使凋落物分解速率减缓,且随氮沉降剂量增加,凋落物分解速率相比对照分别减慢了9.88%（硝态氮低氮）、15.02%（硝态氮高氮）、11.46%（铵态氮低氮）、14.62%（铵态氮高氮）、13.04%（混合态氮低氮）和16.20%（混合态氮高氮）.且不同氮沉降类型、不同氮沉降水平间差异显著.不同形态、不同水平的氮沉降显著地增加了凋落物N 含量（P=0.061,P=0.087）,其中混合态氮沉降对凋落物中N 素含量增加最显著（P=0.044）.但在分解过程中,各处理均未对凋落物C 含量产生显著影响.不同水平的氮沉降显著降低了凋落物的w（C）/w（N）比值,而且不同类型不同水平氮沉降对凋落物w（C）/w（N）比值具有显著的交互作用（P=0.011）.综上所述,通过对模拟氮沉降后凋落物残留率等的变化分析,得出氮沉降对温带森林凋落物的分解产生了抑制作用.     

Decomposition of diverse litter mixtures in streams

In view of growing interest in understanding how biodiversity affects ecosystem functioning, we investigated effects of riparian plant diversity on litter decomposition in forest streams. Leaf litter from 10 deciduous tree species was collected during natural leaf fall at two locations (Massif Central in France and Carpathians in Romania) and exposed in the field in litter bags. There were 35 species combinations, with species richness ranging 1-10. Nonadditive effects on the decomposition of mixed-species litter were minor, although a small synergistic effect was observed in the Massif Central stream where observed litter mass remaining was significantly lower overall than expected from data on single-species litter. In addition, variability in litter mass remaining decreased with litter diversity at both locations. Mean nitrogen concentration of single- and mixed-species litters (0.68-4.47% of litter ash-free dry mass) accounted for a large part of the variation in litter mass loss across species combinations. For a given species or mixture, litter mass loss was also consistently faster in the Massif Central than in the Carpathians, and the similarity in general stream characteristics, other than temperature, suggests that this effect was largely due to differences in thermal regimes. These results support the notion that decomposition of litter mixtures is primarily driven by litter quality and environmental factors, rather than by species richness per se. However, the observed consistent decrease in variability of decomposition rate with increasing plant species richness indicates that conservation of riparian tree diversity is important even when decomposition rates are not greatly influenced by litter mixing.     

Tree species effects on decomposition and forest floor dynamics in a common garden

We studied the effects of tree species on leaf litter decomposition and forest floor dynamics in a common garden experiment of 14 tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Betula pendula, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pinus nigra, Pinus sylvestris, Pseudotsuga menziesii, Quercus robur, Quercus rubra, and Tilia cordata) in southwestern Poland. We used three simultaneous litter bag experiments to tease apart species effects on decomposition via leaf litter chemistry vs. effects on the decomposition environment. Decomposition rates of litter in its plot of origin were negatively correlated with litter lignin and positively correlated with mean annual soil temperature (MAT(soil)) across species. Likewise, decomposition of a common litter type across all plots was positively associated with MAT(soil), and decomposition of litter from all plots in a common plot was negatively related to litter lignin but positively related to litter Ca. Taken together, these results indicate that tree species influenced microbial decomposition primarily via differences in litter lignin (and secondarily, via differences in litter Ca), with high-lignin (and low-Ca) species decomposing most slowly, and by affecting MAT(soil), with warmer plots exhibiting more rapid decomposition. In addition to litter bag experiments, we examined forest floor dynamics in each plot by mass balance, since earthworms were a known component of these forest stands and their access to litter in litter bags was limited. Forest floor removal rates estimated from mass balance were positively related to leaf litter Ca (and unrelated to decay rates obtained using litter bags). Litter Ca, in turn, was positively related to the abundance of earthworms, particularly Lumbricus terrestris. Thus, while species influence microbially mediated decomposition primarily through differences in litter lignin, differences among species in litter Ca are most important in determining species effects on forest floor leaf litter dynamics among these 14 tree species, apparently because of the influence of litter Ca on earthworm activity. The overall influence of these tree species on leaf litter decomposition via effects on both microbial and faunal processing will only become clear when we can quantify the decay dynamics of litter that is translocated belowground by earthworms.     

气候变暖背景下森林土壤碳循环研究进展

由人类活动引起的温室效应以及由此造成的气候变暖对森林牛态系统的影响已引起人们的普遍关注.森林土壤碳循环作为全球碳循环的重要组成部分,是决定未来陆地牛物嘲表现为碳源/碳汇的关键环节,揭示这一作用对于准确理解全球变化背景下陆地生态系统碳循环过程具有重要的指导意义.本文主要通过论述影响土壤碳循环过程的5个方面(土壤呼吸、土壤微生物、土壤酶活性、凋落物输入与分解、土壤碳库),综述了近10 a来全球气候变暖对土壤碳循环过程的影响.近年来,尽管已开展了大量有关土壤碳循环对气候变暖的响应及反馈机制的研究,并取得了一定的成果,但研究结果仍然存在很大的不确定性.整合各种密切关联的全球变化现象,完善研究方法和实验手段,加强根际微生态系统碳循环过程与机理研究将是下一步研究的方向和重点.参70     

Litter decomposition and nitrogen mineralization from an annual to a monthly model

The forest litter decomposition model (FLDM) described in this paper provides an important basis for assessing the impacts of forest management on seasonal stream water quality and export of dissolved organic carbon (DOC). By definition, models with annual time steps are unable to capture seasonal, within-year variation. In order to simulate seasonal variation in litter decomposition and DOC production and export, we have modified an existing annual FLDM to account for monthly dynamics of decomposition and residual mass in experimental litterbags placed in 21 different forests across Canada.The original annual FLDM was formulated with three main litter pools (fast, slow, and very slow decomposing litter) to address the fact that forest litter is naturally composed of a mixture of organic compounds that decompose at different rates. The annual FLDM was shown to provide better simulations than more complex models like CENTURY and SOMM.The revised monthly model retains the original structure of the annual FLDM, but separates litter decomposition from nitrogen (N) mineralization. In the model, monthly soil temperature, soil moisture, and mean January soil temperature are shown to be the most important controlling variables of within-year variation in decomposition. Use of the three variables in a process-based definition of litter decomposition is a significant departure from the empirical definition in the annual model. The revised model is shown to give similar calculations of residual mass and N concentration as the annual model (r² = 0.91, 0.78), despite producing very different timeseries of decomposition over six years. It is shown from a modelling perspective that (i) forest litter decomposition is independent of N mineralization, whereas N mineralization is dependent on litter decomposition, and (ii) mean January soil temperature defines litter decomposition in the summer because of winter-temperatures’ role in modifying forest-floor microorganism community composition and functioning in the following summer.     

保护性耕作对农田土壤有机碳库的影响

农田土壤有机碳库是陆地生态系统碳库的重要组成部分,对维持全球碳平衡具有重要意义。有机碳库的储量除了受气候和环境因素的影响外,还受农业耕作措施的影响。综述了保护性耕作对土壤有机碳库的影响,并从碳输入、碳分解和碳固持3个方面讨论了保护性耕作影响有机碳库的途径。大量实验结果表明,保护性耕作在表层土壤中能提高作物根系生物量1.0%~142.9%;增加土壤微生物的生物量2.2%~140%。免耕还显著降低土壤的呼吸强度,在0.9%~72.6%之间,从而减少碳的损失。免耕还提高了土壤团聚体数量,从而有利于有机碳在土壤中的固持。今后的研究应该在如下几方面加强：（1）加强区域性对比研究;（2）加强长期定位研究;（3）加强对固碳机理的研究;（4）建立模型加强对多种因子综合效应的研究。     

Microbial community composition and function across an arctic tundra landscape

Arctic landscapes are characterized by a diversity of ecosystems, which differ in plant species composition, litter biochemistry, and biogeochemical cycling rates. Tundra ecosystems differing in plant composition should contain compositionally and functionally distinct microbial communities that differentially transform dissolved organic matter as it moves downslope from dry, upland to wet, lowland tundra. To test this idea, we studied soil microbial communities in upland tussock, stream-side birch-willow, and lakeside wet sedge tundra in arctic Alaska, USA. These are a series of ecosystems that differ in topographic position, plant composition, and soil drainage. Phospholipid fatty acid (PLFA) analyses, coupled with compound-specific 13C isotope tracing, were used to quantify microbial community composition and function; we also assayed the activity of extracellular enzymes involved in cellulose, chitin, and lignin degradation. Surface soil from each tundra ecosystem was labeled with 13C-cellobiose,13C-N-acetylglucosamine, or 13C-vanillin. After a five-day incubation, we followed the movement of 13C into bacterial and fungal PLFAs, microbial respiration, dissolved organic carbon, and soil organic matter. Microbial community composition and function were distinct among tundra ecosystems, with tussock tundra containing a significantly greater abundance and activity of soil fungi. Although the majority of 13C-labeled substrates rapidly moved into soil organic matter in all tundra soils (i.e., 50-90% of applied 13C), microbial respiration of labeled substrates in wet sedge tundra soil was lower than in tussock and birch-willow tundra; approximately 8% of 13C-cellobiose and approximately 5% of 13C-vanillin was respired in wet sedge soil vs. 26-38% of 13C-cellobiose and 18-21% of 13C-vanillin in the other tundra ecosystems. Despite these differences, wet sedge tundra exhibited the greatest extracellular enzyme activity. Topographic variation in plant litter biochemistry and soil drainage shape the metabolic capability of soil microbial communities, which, in turn, influence the chemical composition of dissolved organic matter across the arctic tundra landscape.     

The impact of the invasive shrub Lonicera maackii on the decomposition dynamics of a native plant community

Invasive plants may have variable effects within a given environment depending on their interactions with the dominant native species, yet little research has examined such species-species interactions within a site. Savanna trees with nonoverlapping canopies offer an ideal opportunity to assess associated changes in the ecosystem processes that result from interactions between an invasive species and different native tree species. We examined the influence of the exotic invasive shrub Lonicera maackii on decomposition dynamics under three native tree species: Fraxinus quadrangulata, Quercus muehlenbergii, and Carya ovata. Litter decomposition rates and litter C and N were evaluated over two years using single- and mixed-species litterbags (L. maackii and individual tree species litter); microarthropod abundance was measured at 6 weeks using Tulgren funnels. Litter from the invasive L. maackii decomposed and lost N more rapidly than the litter of the three native tree species. The rate at which L. maackii decomposed depended on its location, with L. maackii litter decomposing and losing N more rapidly under C. ovata than under the other two native tree species. Mixing L. maackii with the native species' litter did not accelerate litter mass loss overall but did result in synergistic N losses at variable times throughout the experiment, further highlighting the variable interaction between native species and L. maackii. Nitrogen loss was significantly higher than expected in mixtures of C. ovata + L. maackii litter at 6 weeks, in F. quadrangulata + L. maackii litter at 12 weeks, and in Q. muehlenbergii + L. maackii litter at 24 weeks. If the effects of invasive species on certain ecosystem processes, such as litter decomposition, are strongly influenced by their association with native species, this could suggest the need for a more nuanced understanding of the vulnerability of ecosystem processes to invasions of L. maackii and potentially other invasive species.     

An assessment of seaweed decomposition within a southern Strait of Georgia seaweed community

An assessment of litter and detritus decomposition and nitrogen content of decomposing litter is presented for ten important seaweeds within a southern Strait of Georgia (British Columbia, Canada) seaweed community sampled from August 1975 until October 1976. Litter decomposition rates varied among species with the time required for litter to disappear from litter bags ranging from 6 d for the lamina of Nereocystis luetkeana to about 70 d for Fucus distichus. Decomposition was characterized by an accelerating increase in the nitrogen: dry weight ratio of remnant litter as decomposition proceeded. Iridaea cordata detritus decomposed most rapidly, at 5.7% d^-1, while rates for Gigartina papillata, N. luetkeana, Laminaria saccharina and Laminaria groenlandica ranged from 1.8 to 3.6% d^-1. The remaining species decomposed more slowly. There was a tendency toward more rapid decomposition with decreasing crude fibre content and detritus particle size; however, it appears that morphology, habitat and growth rate are also correlated with relative decomposition rates. Of 43 taxa identified within quantitative litter collections, F. distichus (41%), I. cordata (26%), N. luetkeana (27%) and Laminaria spp. (4%) accounted for 98% of total deposition with mean peak accumulation occurring in August and September from a low near zero in January and February. Litter distribution was patchy, with most litter decomposing near its place of deposition. The application of litter decomposition rates to measured litter accumulation in a mathematical simulation of decomposition predicted the rate of seaweed litter decomposition to peak at about 1.1 g AFDW (ash-free dry weight) m^-2 d^-1 in early September from a mid-winter low near zero. In total, 56±4% of decomposing litter formed detritus, with the remainder being released as soluble matter. The annual contribution of seaweed litter biomass to detrital pathways from our study site was calculated to be 152 g AFDW m^-2.     

Recovery of plant diversity following N cessation: effects of recruitment, litter, and elevated N cycling

Plant species richness has declined and composition shifted in response to elevated atmospheric deposition of biologically active nitrogen over much of the industrialized world. Litter thickness, litter nitrogen (N) content, and soil N mineralization rates often remain elevated long after inputs cease, clouding the prospects that plant community diversity and composition would recover should N inputs be reduced. Here we determined how N cycling, litter accumulation, and recruitment limitation influenced community recovery following cessation of long-term N inputs to prairie-like grasslands. We alleviated each of these potential inhibitors through a two-year full-factorial experiment involving organic carbon addition, litter removal, and seed addition. Seed addition had the largest effect on increasing seedling and species numbers and may be necessary to overcome long-term burial of seeds of target perennial grassland species. Litter removal increased light availability and bare sites for colonization, though it had little effect on reducing the biomass of competing neighbors or altering extractable soil N. Nonetheless, these positive influences were enough to lead to small increases in species richness within one year. We found that, although C addition quickly altered many factors assumed favorable for the target community (decreased N availability and biomass of nearby competitors, increased light and site availability), these changes were insufficient to positively impact species richness or seedling numbers over the experimental duration. However, only carbon addition had species-specific effects on the existing plant community, suggesting that its apparent limited utility may be more a result of slow recovery under ambient recruitment rather than from a lack of a restorative effect. There were dramatic interactions among treatments, with the positive effects of litter removal largely negated by carbon addition, and the positive effects of seed addition generally amplified by litter removal. It remains unclear whether each mechanism explored here will induce community recovery, but over different temporal scales. Long-term monitoring will help resolve these remaining questions. Regardless, our results suggest that reversal of species loss and compositional shifts from N deposition in prairies may be more inhibited by habitat fragmentation, recruitment limitation, and long-term suppression of fire than from continued effects of elevated N.