The role of litter in an old-field community: impact of litter quantity in different seasons on plant species richness and abundance

Summary We studied the effect of removing and adding plant litter in different seasons on biomass, density, and species richness in a Solidago dominated old-field community in New Jersey, USA. We removed all the naturally accumulated plant litter in November (658 g/m²) and in May (856 g/m²) and doubled the amount of litter in November and May in replicated plots (1 m²). An equal number of plots were left as controls. Litter removal and addition had little impact on total plant biomass or individual species biomass in the growing season following the manipulations. Litter removal, however, significantly increased plant densities but this varied depending upon the season of litter removal, species, and life history type. Specifically, the fall litter removal had a much greater impact than the spring litter removal suggesting that litter has its greatest impact after plant senescence in the fall and prior to major periods of early plant growth in spring. Annual species showed the greatest response, especially early in the growing season. Both spring and fall litter removal significantly increased species richness throughout the study. Litter additions in both spring and fall reduced both plant densities and species richness in June, but these differences disappeared near the end of the growing season in September. We concluded than in productive communities where litter accumulation may be substantial, litter may promote low species richness and plant density. This explanation does not invoke resource competition for the decline in species richness. Finally, we hypothesize that there may be broad thresholds of litter accumulation in different community types that may act to either increase or decrease plant yield and diversity.     

Litter accumulation alters the abiotic environment and drives community successional changes in two fenced grasslands in Inner Mongolia

Fencing is an effective and practical method for restoring degraded grasslands in northern China. However, little is known about the role of excess litter accumulation due to long‐term fencing in regulating abiotic environment and driving changes in community structure and function. We conducted a three‐year field experiment in two fenced grasslands in Inner Mongolia, and monitored light quantity, soil temperature, and soil moisture continuously, and determined community height, community aboveground net primary productivity (ANPP), and the relative dominance of different plant functional groups. Litter accumulation reduced light quantity and soil temperature but increased soil moisture. The regulating effects of litter accumulation on soil temperature and soil moisture fluctuated temporally and gradually weakened over the growing season. Litter accumulation also altered community vertical structure and function by increasing community height and ANPP. The increase in soil moisture increased the relative dominance of rhizome grasses but suppressed bunch grasses, thereby shifting bunch grass grasslands to rhizome grass grasslands. Our findings provide a potential mechanism for community succession in the context of litter accumulation in fenced grasslands and indicate that the vegetation and ecosystem services of degraded grasslands are improved after appropriate fencing.     

Can plant litter affect net primary production of a typical steppe in Inner Mongolia?

Question: Litter (dead leaves or stems) affects production by conserving soil moisture. However, that role is not clear for grasslands where most precipitation falls during the growing season when the demand for water is high. Our question was: Does litter affect forage production in such an environment? Location: Typical steppe, Inner Mongolia. Methods: We examined the role of plant litter in two experiments where litter was either removed or added in a protected or heavily grazed site, respectively, in autumn and in spring in a split plot design. The treatments (control, moderate and heavy litter application) were applied once in five replications but repeated at new locations in each of 3 years. This was done to examine only the direct effect of litter on annual net primary production and selected plant characteristics and not potential secondary effects. We also measured soil moisture and soil temperature. Results: Removing litter caused a reduction in the amount of grass (Leymus chinensis) that was produced, but litter addition caused an inconsistent effect among years, with moderate applications producing the most positive effects. Litter removal resulted in shorter and less dense plants of L. chinensis and Carex duriuscula, while heavy litter addition in autumn reduced plant height of both Cleistogenes squarrosa and C. duriuscula. Conclusions: Litter was effective for enhancing soil moisture status and reducing soil heat units in the typical steppe of Inner Mongolia. Therefore, litter mass may serve as an index of grassland health in such environments.     

Altering Rainfall Timing and Quantity in a Mesic Grassland Ecosystem: Design and Performance of Rainfall Manipulation Shelters

Global climate change is predicted to alter growing season rainfall patterns, potentially reducing total amounts of growing season precipitation and redistributing rainfall into fewer but larger individual events. Such changes may affect numerous soil, plant, and ecosystem properties in grasslands and ultimately impact their productivity and biological diversity. Rainout shelters are useful tools for experimental manipulations of rainfall patterns, and permanent fixed-location shelters were established in 1997 to conduct the Rainfall Manipulation Plot study in a mesic tallgrass prairie ecosystem in northeastern Kansas. Twelve 9 x 14–m fixed-location rainfall manipulation shelters were constructed to impose factorial combinations of 30% reduced rainfall quantity and 50% greater interrainfall dry periods on 6 x 6–m plots, to examine how altered rainfall regimes may affect plant species composition, nutrient cycling, and above- and belowground plant growth dynamics. The shelters provided complete control of growing season rainfall patterns, whereas effects on photosynthetic photon flux density, nighttime net radiation, and soil temperature generally were comparable to other similar shelter designs. Soil and plant responses to the first growing season of rainfall manipulations (1998) suggested that the interval between rainfall events may be a primary driver in grassland ecosystem responses to altered rainfall patterns. Aboveground net primary productivity, soil CO₂ flux, and flowering duration were reduced by the increased interrainfall intervals and were mostly unaffected by reduced rainfall quantity. The timing of rainfall events and resulting temporal patterns of soil moisture relative to critical times for microbial activity, biomass accumulation, plant life histories, and other ecological properties may regulate longer-term responses to altered rainfall patterns.     

Invasive grass litter facilitates native shrubs through abiotic effects

Questions: Plant invasions are considered one of the top threats to the biodiversity of native taxa, but clearly documenting the causal links between invasions and the decline of native species remains a major challenge of invasion biology. Most studies have focused on impacts of invaders' living biomass, rather than on mechanisms mediated by litter. However, invasive plant litter, which is often of a very different type and quantity than a system's native plant litter, can have multiple important effects on ecosystem processes – such as nitrogen cycling and soil microclimate – that may influence native plants. Location: We studied effects of litter of invasive grass species that are widespread throughout western North America on native shrubs in southern California's semi‐arid habitat of coastal sage scrub. Methods: We combined a 3‐year field manipulation of non‐native litter with structural equation modeling to understand interacting effects on non‐native grasses, native shrubs, soil nitrogen (available and total), and soil moisture. Results: Litter addition facilitated non‐native grass growth, revealing a positive feedback likely to enhance invasion success. Contrary to a major paradigm of invasion biology – that competition with invasive plant species causes declines of native plants – we found that litter also facilitated growth of the native dominant shrub, a result supported by observational trends. Structural equation models indicated that enhanced soil moisture mediated the positive effects of litter on shrub growth. Conclusions: We demonstrate that invasive plants, via their litter, can facilitate dominant native plants by altering soil moisture. Our results highlight that understanding the impacts and mechanisms of plant invasions may be enhanced by considering the role of invasive plant litter on native plants and ecosystem properties.     

Effect of altered litter input and nitrogen addition on ecosystem aboveground primary productivity and plant functional group composition in a semiarid grassland

不同的草原利用方式(围封、放牧和割草等)随着大气氮沉降的不断加剧, 改变了凋落物输入量。凋落物作为连接地上-地下碳循环过程的关键环节, 对草原生态系统生产力和碳循环过程影响显著。氮是草原生产力的主要限制因子, 凋落物输入量的变化对草原生态系统结构和功能的影响仍缺乏长期实验证据支持。该研究在内蒙古半干旱典型草原建立一个凋落物输入变化和氮添加控制实验平台, 通过连续6年对群落生产力和功能群组成的监测, 研究了凋落物添加与去除和氮添加对半干旱草原群落生产力和功能群组成的影响。研究发现: 1)凋落物输入量增加和氮添加均显著提高了群落生产力, 在对照和氮添加处理下, 凋落物去除处理导致生产力分别降低了8.4%和7.6%, 而凋落物添加处理使生产力分别提高了10.7%和6.3%; 2)不同植物功能群对凋落物输入变化和氮添加的响应存在差异, 导致群落功能群结构发生变化。随着凋落物输入量增加和氮添加, 群落优势功能群多年生禾草(包括多年生丛生禾草和多年生根茎禾草)的生物量显著提高, 对群落生产力的贡献增加, 在群落中的优势地位增强; 而另一优势功能群多年生杂类草生物量对凋落物和氮添加处理均无显著响应, 进而导致在氮添加处理下其对群落生物量的贡献比例显著降低; 3)凋落物输入主要改善土壤水分状况, 而氮添加则主要通过提高土壤养分含量, 促进群落生产力, 并通过影响主要功能群生物量, 导致群落结构发生变化。以上结果表明, 适当的草原管理方式如围封禁牧和降低放牧强度等都能通过增加凋落物的输入来提高草原生产力, 维持生态系统稳定性。而适量的氮等养分添加管理也有助于提高草原生产力, 促进其恢复。     

Nitrogen addition overrides the effects of Stellera chamaejasme litter on the growth of Leymus chinensis and its associated mycorrhizal fungi

氮添加掩盖了狼毒凋落物对羊草生长及其菌根真菌的影响 退化草地上迅速蔓延、生长的有毒植物能产生大量的凋落物,而氮输入能促进有毒植物的生长, 并改变其根际微生物特别是丛枝菌根真菌(AMF)的特征。然而,有毒植物的凋落物如何影响可食牧草的生长及其根际AMF,N沉降是否以及如何介导这些影响,目前尚不清楚。本文在中国东北的草甸草原上进行了温室实验以检验添加狼毒(Stellera chamaejasme)凋落物和氮素对优势植物羊草(Leymus chinensis) 的生长、AMF和土壤特征的影响。结果表明,添加凋落物显著增加了羊草分株数、地上生物量和土壤速效磷(AP)浓度,并降低了AMF孢子密度。然而,凋落物与氮素的交互作用并未显著影响羊草和AMF。狼毒凋落物通过提高AP对羊草产生正效应,并结合土壤养分平衡与凋落物诱导的化感成分的变化对AMF产生负效应。另外,高氮添加能缓解土壤的氮限制并抑制凋落物分解,从而减缓凋落物对羊草和AMF的影响。上述结果表明,除了关注有毒植物对家畜的有害作用外,还应客观地、全面地评价有毒植物的生态功能;同时,氮沉降应作为预测退化草地有毒植物与可食植物的种间关系的一项重要指标。     

不同月份采集的贝加尔针茅地上部的分解特征及影响因素

利用植物生长过程中植物组织内养分元素化学性质的季节性变化特点,在2009年生长季的不同月份(5—9月)采集了呼伦贝尔地区贝加尔针茅地上部(茎和叶),作为分解底物,采用分解袋法,研究其分解特点及其影响因素。结果表明:不同月份采集的贝加尔针茅分解底物之间的分解速度存在明显差异,依采集月份逐渐递减,5月>6月>7月>8月>9月;分解底物初始N、P、Zn、K、Mg和Mn含量与分解速度均呈显著正相关,而碳含量和C∶N与分解速度呈显著负相关,初始钙含量、N∶P与分解速度无显著相关性;土壤微生物生物量N与分解底物的残余质量(%)相关性明显;另外,土壤微生物生物量N与凋落物C、N含量也具有明显的相关性;分解底物质量损失与土壤水分含量显著正相关,而与土壤温度的相关性较弱,说明降水变化通过调节土壤湿度来影响凋落物分解。     

凋落物输入变化和氮添加对半干旱草原群落生产力及功能群组成的影响

不同的草原利用方式(围封、放牧和割草等)随着大气氮沉降的不断加剧, 改变了凋落物输入量。凋落物作为连接地上-地下碳循环过程的关键环节, 对草原生态系统生产力和碳循环过程影响显著。氮是草原生产力的主要限制因子, 凋落物输入量的变化对草原生态系统结构和功能的影响仍缺乏长期实验证据支持。该研究在内蒙古半干旱典型草原建立一个凋落物输入变化和氮添加控制实验平台, 通过连续6年对群落生产力和功能群组成的监测, 研究了凋落物添加与去除和氮添加对半干旱草原群落生产力和功能群组成的影响。研究发现: 1)凋落物输入量增加和氮添加均显著提高了群落生产力, 在对照和氮添加处理下, 凋落物去除处理导致生产力分别降低了8.4%和7.6%, 而凋落物添加处理使生产力分别提高了10.7%和6.3%; 2)不同植物功能群对凋落物输入变化和氮添加的响应存在差异, 导致群落功能群结构发生变化。随着凋落物输入量增加和氮添加, 群落优势功能群多年生禾草(包括多年生丛生禾草和多年生根茎禾草)的生物量显著提高, 对群落生产力的贡献增加, 在群落中的优势地位增强; 而另一优势功能群多年生杂类草生物量对凋落物和氮添加处理均无显著响应, 进而导致在氮添加处理下其对群落生物量的贡献比例显著降低; 3)凋落物输入主要改善土壤水分状况, 而氮添加则主要通过提高土壤养分含量, 促进群落生产力, 并通过影响主要功能群生物量, 导致群落结构发生变化。以上结果表明, 适当的草原管理方式如围封禁牧和降低放牧强度等都能通过增加凋落物的输入来提高草原生产力, 维持生态系统稳定性。而适量的氮等养分添加管理也有助于提高草原生产力, 促进其恢复。     

Ecological significance of litter redistribution by wind and snow in arctic landscapes

During winter in the Arctic, plant litter is scoured from exposed hills and ridges by wind and snow and is redistributed to other portions of the landscape. The aim of this research was to quantify the physical and biological consequences of this litter redistribution. Litter biomass accumulation was ten times greater in areas of high deposition (e.g. snow drifts) than in areas of low deposition. Spring snow melt was delayed by several days and soils were cooler throughout the growing season and throughout winter in areas of increased litter deposition than areas with no litter. Photosynthetically active radiation (PAR) was reduced to near zero with small accumulations of litter. Annual C and N inputs from allochthonous litter were 143 g Cm-2 and 14 g Nm in high lilter areas and 3.4 g C m⁻² and 0.3 g N ^m-2 in non-drift, ambient litter deposition areas. Although PAR and soil temperatures were significantly reduced with increased litter deposition, we did not observe significant delays in key plant phenological events of several species or measure a decrease in gross ecosystem photosynthesis. We did measure a significant increase in ecosystem respiration with increased litter deposition, which resulted in a shift in the net C balance of dry heath tundra from near zero with no litter to a net source of CO₂ to the atmosphere. Our study indicates that the redistribution of litter by wind and snow during winter is an important mechanism of nutrient transfer across the arctic landscape and that allochthonous litter inputs are of great enough magnitude to alter the carbon balance of some areas of the arctic landscape.     

短期增温和增加降水对内蒙古荒漠草原土壤呼吸的影响

利用红外辐射增温装置模拟短期持续增温和降水增加交互作用对内蒙古荒漠草原土壤呼吸作用的影响, 结果表明: 土壤含水量对月土壤呼吸的影响显著大于土壤温度增加的影响, 生长旺季的月土壤呼吸显著大于生长末季; 土壤温度和水分增加都显著影响日土壤呼吸, 但二者的交互作用对土壤呼吸无显著影响。荒漠草原7‒8月平均土壤呼吸速率为1.35 μmol CO₂·m ^-2·s ^-1, 7月份为2.08 μmol CO₂·m ^-2·s ^-1, 8月份为0.63 μmol CO₂·m ^-2·s ^-1。土壤呼吸与地下各层根系生物量呈幂函数关系, 0‒10 cm土层的根系生物量对土壤呼吸的解释率(79.2%)明显高于10‒20 cm土层的解释率(31.6%)。0-10 cm土层的根系生物量是根系生物量的主体, 根系生物量对土壤呼吸的影响具有层次性。在未来全球变暖和降水格局变化的情景下, 荒漠草原土壤水分含量是影响生物量的主导环境因子, 而根系生物量的差异是造成土壤呼吸异质性的主要生物因素, 土壤含水量可通过影响根系生物量控制土壤呼吸的异质性。     

A water-centred framework to assess the effects of salinity on the growth and yield of wheat and barley

We conducted a field experiment in two alpine meadows to investigate the short-term effects of nitrogen enrichment and plant litter biomass on plant species richness, the percent cover of functional groups, soil microbial biomass, and enzyme activity in two alpine meadow communities. The addition of nitrogen fertilizer to experimental plots over two growing seasons increased plant production, as indicated by increases in both the living plant biomass and litter biomass in the Kobresia humilis meadow community. In contrast, fertilization had no significant effect on the amounts of living biomass and litter biomass in the K. tibetica meadow. The litter treatment results indicate that litter removal significantly increased the living biomass and decreased the litter biomass in the K. humilis meadow; however, litter-removal and litter-intact treatments had no impact on the amounts of living biomass and litter biomass in the K. tibetica meadow. Litter production depended on the degree of grass cover and was also influenced by nitrogen enrichment. The increase in plant biomass reflects a strong positive effect of nitrogen enrichment and litter removal on grasses in the K. humilis meadow. Neither fertilization nor litter removal had any impact on the grass biomass in the K. tibetica meadow. Sedge biomass was not significantly affected by either nutrient enrichment or litter removal in either alpine meadow community. The plant species richness decreased in the K. humilis meadow following nitrogen addition. In the K. humilis meadow, microbial biomass C increased significantly in response to the nitrogen enrichment and litter removal treatments. Enzyme activities differed depending on the enzyme and the different alpine meadow communities; in general, enzyme activities were higher in the upper soil layers (0–10 cm and 10–20 cm) than in the lower soil layers (20–40 cm). The amounts of living plant biomass and plant litter biomass in response to the different treatments of the two alpine meadow communities affected the soil microbial biomass C, soil organic C, and soil fertility. These results suggest that the original soil conditions, plant community composition, and community productivity are very important in regulating plant community productivity and microbial biomass and activity.     

Subordinate plant species impact on soil microbial communities and ecosystem functioning in grasslands: Findings from a removal experiment

Despite their low relative abundance, subordinate plant species may have larger impacts on ecosystem functioning than expected, but their role in plant communities remains poorly understood. The aim of this study was to test how subordinate plant species influence the functioning of a species-rich semi-natural grasslands. A plant removal experiment was set-up in the mountain grasslands of the Jura Mountains (Switzerland) to test the impact of subordinate plant species on soil microbial communities and ecosystem functioning. The experiment included three treatments: removal of all subordinate species, partial biomass removal of dominant species, and a no biomass removal control. After 2 years of treatments, we determined soil microbial community (bacteria and mycorrhizal fungi) by T-RFLP analysis and measured litter decomposition, soil respiration, soil inorganic nitrogen (DIN) availability and throughout above-ground biomass production as measures of ecosystem function. The removal of subordinate plant species strongly affected bacterial and weakly influenced mycorrhizal fungi communities and decreased rates of plant litter decomposition, soil respiration and DIN availability with larger effects than the partial loss of dominant biomass. The removal of subordinate plant species did not modify plant community structure, but it did reduce total above-ground biomass production compared to the control plots. Collectively, our findings indicate that the loss of subordinate species can have significant consequences for soil microbial communities and ecosystem functions, suggesting that subordinate species are important drivers of ecosystem properties.     

典型草原围封后羊草地上功能性状对枯落物累积的响应

植物功能性状反映植物适应环境变化过程中在不同器官形态与功能间的资源权衡与分配策略。典型草原围封后枯落物累积导致群落光照、热量和水分的重新分配并改变微环境特征。在此过程中植物地上功能性状将通过怎样的变化来适应新的环境,目前尚不清楚。2015—2017年每年8月对内蒙古地区3种典型草原共有物种羊草（Leymus chinensis）的植株、叶片和茎干功能性状进行了测量与分析。结果表明：枯落物累积显著增加了羊草的植株高度、单株重量、茎叶比和总叶面积;枯落物累积显著增加了羊草的叶片长度、叶片重量、单叶面积、节间长度和茎干重量,这些性状属于敏感性状;枯落物累积对羊草的叶片数量和节间数量无显著影响,相对而言,这些性状属于惰性性状;羊草的单株重量与植株高度、叶片重量呈极显著的正相关关系（P<0.0001）;羊草的植株高度与节间数量、节间长度呈极显著的正相关关系（P<0.0001）。本研究结果从植物地上功能性状的角度阐明了典型草原植物对环境变化的适应方式,可为围封草原的合理管理提供基础数据与理论依据。     

Carbon dioxide fluxes in a spatially and temporally heterogeneous temperate grassland

Landscape position, grazing, and seasonal variation in precipitation and temperature create spatial and temporal variability in soil processes, and plant biomass and composition in grasslands. However, it is unclear how this variation in plant and soil properties affects carbon dioxide (CO₂) fluxes. The aim of this study is to explore the effect of grazing, topographic position, and seasonal variation in soil moisture and temperature on plant assimilation, shoot and soil respiration, and net ecosystem CO₂ exchange (NEE). Carbon dioxide fluxes, vegetation, and environmental variables were measured once a month inside and outside long-term ungulate exclosures in hilltop (dry) to slope bottom (mesic) grassland throughout the 2004 growing season in Yellowstone National Park. There was no difference in vegetation properties and CO₂ fluxes between the grazed and the ungrazed sites. The spatial and temporal variability in CO₂ fluxes were related to differences in aboveground biomass and total shoot nitrogen content, which were both related to variability in soil moisture. All sites were CO₂ sinks (NEE>0) for all our measurments taken throughout the growing season; but CO₂ fluxes were four- to fivefold higher at sites supporting the most aboveground biomass located at slope bottoms, compared to the sites with low biomass located at hilltops or slopes. The dry sites assimilated more CO₂ per gram aboveground biomass and stored proportionally more of the gross-assimilated CO₂ in the soil, compared to wet sites. These results indicate large spatio-temporal variability of CO₂ fluxes and suggest factors that control the variability in Yellowstone National Park.     

草地利用方式对土壤呼吸和凋落物分解的影响

草地利用方式影响植被群落结构和土壤微环境, 制约草地生态系统碳循环。该文通过测定温带草原在放牧、割草、围封3种利用方式下湿润年(2012年)和干旱年(2011年)的凋落物产量、质量及其分解速率和土壤碳通量, 分析了草地利用方式对土壤呼吸和凋落物的影响, 探讨了凋落物对土壤呼吸的贡献机制。结果表明: 在干旱年份, 放牧样地土壤呼吸最大, 分别达到割草和围封样地的1.5倍和1.29倍; 在湿润年份, 割草样地土壤呼吸最大, 为309 g C∙m^-2∙a^-1, 明显高于放牧样地和围封样地。不论干旱年还是湿润年, 围封样地凋落物产量都大于放牧样地和割草样地。3种利用方式下湿润年土壤呼吸和凋落物分解均比干旱年增强。因此, 水分是温带草原植物生长和生态系统碳循环的主要限制因子, 草地利用方式则显著影响凋落物生产和分解。进一步分析表明, 经过两年的分解, 同一样地内凋落物质量C:N下降, N含量和木质素:N升高, 土壤呼吸与凋落物产量、凋落物分解速率以及木质素:N正相关, 而与凋落物C:N负相关。     

Plant–soil feedback during biological invasions: effect of litter decomposition from an invasive plant (Sphagneticola trilobata) on its native congener (S. calendulacea)

生物入侵过程中的植物-土壤反馈：一种入侵植物的凋落物分解对其本地近缘植物的影响 植物入侵可通过正或负的植物-土壤反馈效应改变土壤的生物和非生物性质,从而影响入侵栖息地的土壤理化性质。许多入侵物种的凋落物分解可增加土壤养分,降低本地植物多样性,并导致进一步的植物入侵。关于入侵植物凋落物在不同土壤类型及深度分解及反馈效应的研究依然很少。本研究旨在明确入侵植物南美蟛蜞菊(Sphagneticola trilobata)凋落物在不同土壤类型和不同土壤深度条件下的分解情 况及其对本地近缘植物蟛蜞菊(S. calendulacea)生理生长的影响。将装有南美蟛蜞菊凋落物的尼龙袋加入到不同深度(即0、2、4 和6 cm)的砂土、营养土和粘土中,经6个月的分解后,回收凋落物袋并计算分解速率,随后在凋落物分解处理后的土壤中种植本地蟛蜞菊,并在生长期结束时测量其生理生态指标。研究结果表明,所有处理土壤类型中,凋落物在土壤深度为2和4 cm处分解后显著增加了土壤养分,而对本 地蟛蜞菊的叶片叶绿素、叶氮含量等生长指标表现为负效应。因此,入侵植物南美蟛蜞菊凋落物分解对土壤养分表现为正的反馈效应,而对本地植物蟛蜞菊的生长表现为负效应。我们的研究结果还表明,入侵植物的凋落物分解对土壤和本地物种的影响还因凋落物分解所在的土壤深度而显著不同。未来的研究应侧重于入侵栖息地中更多本地和入侵物种的植物-土壤反馈效应,以及更多土壤类型和土壤深度的入侵植物凋落物效应。     

Plant litter: Its dynamics and effects on plant community structure

We discuss the dynamics of plant litter, the effects of litter on the chemical and physical environment, the direct and indirect effects of plant litter on plant populations and communities, and different adaptative traits that may be related to litter accumulation. The production of litter depends primarily on the site productivity, but other properties of the environment, as well as chance, may introduce important variation. The existence of time lags between the production of plant organs and their transformation into litter appears as a relevant character of litter dynamics seldom included in models. Herbivory, and other processes that destroy biomass or reduce productivity, may reduce the amount of litter produced. The destruction of litter encompasses a complex of interactions. The main processes, including physical and chemical degradation, consumption by invertebrates and decomposition, are differentially affected by the environment and by the physical and chemical characteristics of the litter itself. The relative importance of those processes varies among systems. Litter alters the physical and chemical environment directly and indirectly. The decomposition of litter may release both nutrients and phytotoxic substances into the soil. The physical changes produced by litter also alter the activity of decomposers, resulting in an indirect effect on the chemical environment. The accumulated litter intercepts light, shading seeds and seedlings, and reduces the thermal amplitude in the soil. By reducing maximum soil temperatures, and creating a barrier to water vapor diffusion, litter reduces evaporation from the soil. However, litter may also diminish water availability when it retains a large proportion of rainfall. Litter creates a physical barrier for seedling and sprout emergence and to seeds reaching the soil.     

Cessation of Burning Dries Soils Long Term in a Tallgrass Prairie

Soil moisture is a critical variable in grassland function, yet how fire regimes influence ecohydrology is poorly understood. By altering productivity, species composition, and litter accumulation, fire can indirectly increase or decrease soil water depletion on a range of time scales and depths in the soil profile. To better understand how fire influences soil moisture in grasslands, we analyzed 28 years of soil moisture data from two watersheds in a central North American grassland which differ in their long-term fire frequency. Across 28 years, cessation of prescribed burning initially led to wetter soils, likely as litter accumulated and both transpiration and evaporation were suppressed. Long-term, cessation of burning led to soils drying more, especially at depths greater than 75 cm. The long-term drying of deep soils is consistent with the increase in woody species in the infrequently burned grassland as woody species likely have a greater reliance on soil water from deeper soil layers compared to co-occurring herbaceous species. Despite the ecohydrological changes associated with the cessation of prescribed burning, watersheds with different burn regimes responded similarly to short-term variation in climate variation. In both watersheds, low precipitation and high temperatures led to drier soils with greater responses in soil moisture to climate variation later in the season than earlier. There is no current evidence that the cessation of burning in this ecosystem will qualitatively alter how evapotranspiration responds to climate variation, but the use of deeper soil water by woody plants has the potential for greater transpiration during dry times. In all, modeling the depth-specific responses of soil moisture and associated ecosystem processes to changes in burn regimes will likely require including responses of plant community composition over short and long time scales.     

Asynchronous fluctuation of soil microbial biomass and plant litterfall in a tropical wet forest

Carbon availability often controls soil microbial growth and there is evidence that at regional scales soil microbial biomass is positively correlated with aboveground forest litter input. We examined the influence of plant litterfall on annual variation of soil microbial biomass in control and litter-excluded plots in a tropical wet forest of Puerto Rico. We also measured soil moisture, soil temperature, and plant litterfall in these treatment plots. Aboveground plant litter input had no effect on soil microbial biomass or on its pattern of fluctuation. Monthly changes in soil microbial biomass were not synchronized with aboveground litter inputs, but rather preceeded litterfall by one month. Soil microbial biomass did not correlate with soil temperature, moisture, or rainfall. Our results suggest that changes in soil microbial biomass are not directly regulated by soil temperature, moisture, or aboveground litter input at local scales within a tropical wet forest, and there were asynchronous fluctuation between soil microbial biomass and plant litterfall. Potential mechanisms for this asynchronous fluctuation include soil microbial biomass regulation by competition for soil nutrients between microorganisms and plants, and regulation by below-ground carbon inputs associated with the annual solar and drying-rewetting cycles in tropical wet forests.