Invasion by <Emphasis Type="Italic">Aegilops triuncialis</Emphasis> (Barb Goatgrass) Slows Carbon and Nutrient Cycling in a Serpentine Grassland

Invasive plant species alter plant community composition and ecosystem function. In the United States, California native grasslands have been displaced almost completely by invasive annual grasses, with serpentine grasslands being one of the few remaining refugia for California grasslands. This study examined how the invasive annual grass, Aegilops triuncialis, has altered decomposition processes in a serpentine annual grassland. Our objectives were to (1) assess howA. triuncialis alters primary productivity and litter tissue chemistry, (2) determine whether A. triuncialis litter is more recalcitrant to decomposition than native litter, and (3) evaluate whether differences in the soil microbial community in A. triuncialis-invaded and native-dominated areas result in different decomposition rates of invasive and/or native plant litter. In invaded plant patches, A. triuncialis was approximately 50% of the total plant cover, in contrast to native plant patches in which A. triuncialis was not detected and native plants comprised over 90% of the total plant cover. End-of-season aboveground biomass was 2-fold higher in A. triuncialis dominated plots compared to native plots; however, there was no significant difference in belowground biomass. Both above- and below-ground plant litter from A. triuncialis plots had significantly higher lignin:N and C:N ratios and lower total N, P, and K than litter from native plant plots. Aboveground litter from native plots decomposed more rapidly than litter from A. triuncialis plots, although there was no difference in decomposition of belowground tissues. Soil microbial community composition associated with different soil patch types had no effect on decomposition rates. These data suggest that plant invasion impacts decomposition and nutrient cycling through changes in plant community tissue chemistry and biomass production.     

Grass invasion causes rapid increases in ecosystem carbon and nitrogen storage in a semiarid shrubland

Accurately predicting terrestrial carbon (C) and nitrogen (N) storage requires understanding how plant invasions alter cycling and storage. A common, highly successful type of plant invasion occurs when the invasive species is of a distinctly different functional type than the native dominant plant, such as shrub encroachment throughout the western United States and annual grass invasions in Mediterranean shrublands, as studied here. Such invasions can dramatically transform landscapes and have large potential to alter C and N cycling by influencing storage in multiple pools. We used a manipulation of non‐native annual grass litter within a shrub‐dominated habitat in southern California (coastal sage scrub, CSS) to study how grass invasion alters ecosystem C and N storage. We added, removed, or left unchanged grass litter in areas of high and low invasion, then followed soil and vegetation changes. Grass litter greatly increased C and N storage in soil, aboveground native and non‐native biomass. Aboveground litter storage increased due to the greater inputs and slower decomposition of grass litter relative to shrub litter; shading by grass litter further reduced decomposition of both non‐native and native litter, which may be due to reduced photodegradation. Soil C and N pools in areas of high litter increased ～20% relative to low litter areas in the two years following manipulation and were generally sinks for C and N, while areas with low litter were sources. We synthesize our results into a C cycle of invaded and uninvaded areas of CSS and link changes in storage to increases in the soil fungi : bacteria ratio, increased plant inputs, and decreased litter loss. Overall, we show that grasses, especially through their litter, control important abiotic and biotic mechanisms governing C and N storage, with widespread implications for C sequestration and N storage in semiarid systems undergoing grass or shrub invasions.     

降水变化和氮沉降对荒漠草原两种多年生禾草凋落物分解的影响

凋落物分解是陆地生态系统养分循环的重要过程,在生物地球化学循环过程中发挥着重要作用。全球变化是影响凋落物分解的重要因子,其对生态系统养分循环的影响存在诸多不确定性。研究荒漠草原凋落物分解对氮沉降和降水变化及其二者交互作用的响应,是揭示这些不确定性、保护草原生态系统结构和功能的科学基础。以内蒙古四子王旗短花针茅荒漠草原为研究对象,选取建群种短花针茅和优势种无芒隐子草两种植物凋落物,开展为期4年的长期分解实验,探究两种植物凋落物分解特征及养分释放规律。实验采用裂区设计,主区为自然降水（C）、增雨30%（W）和减雨30%（R）3个水分梯度,副区为0（N0）、30（N30）、50（N50）和100（N100） kg hm^-2 a^-1 4个氮素梯度。结果表明：（1）增雨和氮沉降促进荒漠草原凋落物分解,减雨反之,降水对两种凋落物影响具有差异,而氮沉降的作用不依赖于物种;（2）氮沉降缩短凋落物分解周期5.12%-14.82%,增雨与氮沉降交互缩短凋落物分解周期3.69%-28.75%;（3）降水始终有利于凋落物中碳、纤维素和木质素释放,而分解后期氮沉降对其影响不显著,凋落物分解后期主要受木质素分解速率控制。综上所述,影响荒漠草原凋落物分解的主要因素为降水,其次是氮素,二者对凋落物分解具有协同作用。     

Decomposition of litter produced under elevated CO2: Dependence on plant species and nutrient supply

We investigated the effect of CO₂ concentration and soilnutrient availability during growth on the subsequent decomposition andnitrogen (N) release from litter of four annual grasses that differ inresource requirements and native habitat. Vulpia microstachys isa native grass found on California serpentine soils, whereas Avenafatua, Bromus hordaceus, and Lolium multiflorum areintroduced grasses restricted to more fertile sandstone soils (Hobbs & Mooney 1991). Growth in elevated CO₂ altered litter C:N ratio,decomposition, and N release, but the direction and magnitude of thechanges differed among plant species and nutrient treatments. ElevatedCO₂ had relatively modest effects on C:N ratio of litter,increasing this ratio in Lolium roots (and shoots at high nutrients),but decreasing C:N ratio in Avena shoots. Growth of plants underelevated CO₂ decreased the decomposition rate of Vulpialitter, but increased decomposition of Avena litter from the high-nutrient treatment. The impact of elevated CO₂ on N loss fromlitter also differed among species, with Vulpia litter from high-CO₂ plants releasing N more slowly than ambient-CO₂litter, whereas growth under elevated CO₂ caused increased Nloss from Avena litter. CO₂ effects on N release in Lolium and Bromus depended on the nutrient regime in whichplants were grown. There was no overall relationship between litter C:Nratio and decomposition rate or N release across species and treatments.Based on our study and the literature, we conclude that the effects ofelevated CO₂ on decomposition and N release from litter arehighly species-specific. These results do not support the hypothesis thatCO₂ effects on litter quality consistently lead to decreasednutrient availability in nutrient-limited ecosystems exposed to elevatedCO₂.     

凋落物多样性及组成对凋落物分解及土壤微生物群落的影响——二氧化碳倍增条件下

受全球变化的影响生物多样性的丧失日益严重,及时了解凋落物物种多样性及其组成如何直接或者通过调节微生物群落来间接影响凋落物分解已经成为生态学领域的热点问题之一。在呼伦贝尔典型草原区,羊草（Leymus chinensis）为原生群落建群种,茵陈蒿（Artemisia capillaris）、麻花头（Serratula centauroides）、二裂委陵菜（Potentilla bifurca）在退化及恢复群落中的多度均较大,本研究以此4种植物的凋落物为研究对象,在两倍于当前大气CO₂浓度（800 μmol/mol）条件下,通过嵌套实验设计来研究凋落物多样性（凋落物组成）对凋落物质量、C、N残余率和残余C/N的影响,并深入分析凋落物初始性质以及土壤革兰氏阳性菌（G⁺）、革兰氏阴性菌（G^-）、细菌（B）、真菌（F）及土壤总微生物磷脂脂肪酸（Phospholipid Fatty Acid,PLFA）含量和F/B对凋落物分解的影响。结果表明：（1）凋落物多样性及组成对凋落物质量、C、N残余率以及残余C/N均具有显著影响;凋落物组成对G⁺ PLFAs含量具有显著影响;（2）冗余分析（Redundancy Analysis,RDA）结果表明凋落物组成对凋落物分解和土壤微生物群落相关指标的影响高于凋落物多样性;（3）结构方程模型（Structural Equation Model,SEM）结果表明凋落物初始木质素含量和初始C/N均对凋落物分解具有显著影响,其中凋落物初始木质素含量起主导作用,其还可通过对土壤真菌PLFAs含量的影响来间接影响凋落物N残余率和残余C/N。所得结果可为大气CO₂浓度升高条件下退化草原生态系统的物质循环特征的预测乃至草原生态系统功能的合理评估提供数据支持。     

The impact of elevated CO2, increased nitrogen availability and biodiversity on plant tissue quality and decomposition

Elevated CO₂, increased nitrogen (N) deposition and increasing species richness can increase net primary productivity (NPP). However, unless there are comparable changes in decomposition, increases in productivity will most likely be unsustainable. Without comparable increases in decomposition nutrients would accumulate in dead organic matter leading to nutrient limitations that could eventually prohibit additional increases in productivity. To address this issue, we measured aboveground plant and litter quality and belowground root quality, as well as decomposition of aboveground litter for one and 2‐year periods using in situ litterbags in response to a three‐way factorial manipulation of CO₂ (ambient vs. 560 ppm), N deposition (ambient vs. the addition of 4 g N m⁻² yr⁻¹) and plant species richness (one, four, nine and 16 species) in experimental grassland plots. Litter chemistry responded to the CO₂, N and plant diversity treatments, but decomposition was much less responsive. Elevated CO₂ induced decreases in % N and % lignin in plant tissues. N addition led to increases in % N and decreases in % lignin. Increasing plant diversity led to decreases in % N and % lignin and an increase in % cellulose. In contrast to the litter chemistry changes, elevated CO₂ had a much lower impact on decomposition and resulted in only a 2.5% decrease in carbon (C) loss. Detectable responses were not observed either to N addition or to species richness. These results suggest that global change factors such as biodiversity loss, elevated CO₂ and N deposition lead to significant changes in tissue quality; however, the response of decomposition is modest. Thus, the observed increases in productivity at higher diversity levels and with elevated CO₂ and N fertilization are not matched by an increase in decomposition rates. This lack of coupled responses between production and decomposition is likely to result in an accumulation of nutrients in the litter pool which will dampen the response of NPP to these factors over time.     

不同土壤厚度、水分和种植方式对喀斯特两种草本凋落物分解质量损失和化学计量特征的影响

为了探究生长期间不同土壤厚度、水分及种植方式处理对草本植物凋落物分解质量损失和化学计量特征的的影响,采用分解袋法,在露天分解床上分解经过生长期间2种土壤厚度(对照土壤厚度和浅土处理)、2种水分(正常灌水处理和干旱处理)和2种种植方式(单种和混种)处理的苇状羊茅(Festuca arundinacea Schreb.)和黑麦草(Lolium perenne L.)凋落物,研究生长期间土壤厚度和水分减少及不同的种植方式是否通过改变两物种初始凋落物质量、产量和组分来影响自身凋落物分解。结果发现:(1)与对照组(CK)相比,在干旱组(D)和浅土+干旱组(SD),两物种地上、根系和总的凋落物质量损失率、初始N和P含量均显著增加,凋落物产量、C/N和C/P显著降低,凋落物地上组分比大体上无显著变化;而不同的种植方式处理对各组分质量损失率、元素含量、计量比、凋落物产量和组分比大体上无显著影响;(2)两物种地上、根系和总凋落物的质量损失率分别与地上、根系和总N含量呈显著正相关,与C/N呈显著负相关,与凋落量呈显著负相关,而总凋落物质量损失率与地上凋落物组分比呈显著正相关。结果表明,生长期间干旱和浅土+干旱处理能够通过影响苇状羊茅和黑麦草的初始凋落物质量、产量和组分比来加快地上、根系和总凋落物分解,其中凋落物N含量和C/N是影响两物种凋落物分解快慢的主要原因。     

凋落物多样性及组成对凋落物分解和土壤微生物群落的影响

凋落物分解是生态系统营养物质循环的核心过程,而土壤微生物群落在凋落物分解过程中扮演着极其重要且不可替代的角色。随着生物多样性的丧失日益严峻,探讨凋落物多样性及组成对凋落物分解和土壤微生物群落的影响,不仅有助于了解凋落物分解的内在机制,而且可为退化草原生态系统的恢复提供参考。以内蒙古呼伦贝尔草原退化恢复群落中的草本植物为研究对象,依据植物多度、盖度、频度和物种的重要值及其在群落中的恢复程度筛选出排序前4的羊草（Leymus chinensis）、茵陈蒿（Artemisia capillaris）、麻花头（Serratula centauroides）、二裂委陵菜（Potentilla bifurca）的凋落物为实验材料,通过设置3种凋落物多样性水平（1,2,4）,包括11种凋落物组合（单物种凋落物共4种,两物种凋落物混合共6种,四物种凋落物混合共1种）,利用磷脂脂肪酸（PLFA）方法来研究分解60 d后凋落物多样性及组成对凋落物分解和土壤微生物群落的影响。结果表明：（1）凋落物物种多样性仅对C残余率具有显著影响,表现在两物种混合凋落物C残余率显著低于单物种凋落物,而凋落物组成对所观测的4个凋落物分解参数（质量、C、N残余率以及C/N）均具有显著影响;（2）凋落物物种多样性对细菌（B）含量具有显著影响,而凋落物组成对真菌（F）含量具有显著影响,两者对F/B以及微生物总量均无显著影响;（3）冗余分析结果表明凋落物组成与凋落物分解相关指标（凋落物质量、C、N残余率及C/N）和土壤微生物（真菌、细菌含量）的相关关系高于凋落物多样性。（4）进一步建立结构方程模型（Structural Equation Model,SEM）发现,凋落物初始C含量对凋落物质量、C、N残余率及C/N有显著正的直接影响;凋落物木质素含量对凋落物质量、C、N残余率有显著正的直接影响;凋落物初始N含量对N残余率有显著正的直接影响,而对C残余率及C/N有显著负的直接影响;凋落物初始C/N对凋落物质量、N残余率有显著正的直接影响,而对C/N有显著负的直接影响。此外,凋落物初始C、N、木质素含量及C/N均对真菌含量具有显著正影响,并可通过真菌对凋落物质量分解产生显著负的间接影响。以上结果表明该退化恢复区域优势种凋落物分解以初始C、木质素为主导,主要通过土壤真菌影响凋落物的分解进程,这将减缓凋落物的分解速率进而减慢草原生态系统的进程。这些结果为凋落物多样性及组成对自身分解和土壤微生物群落的影响提供了实验依据,也为进一步分析凋落物分解内在机制以及草原生态系统的恢复提供了数据参考。     

Plant litter effects on soil nutrient availability and vegetation dynamics: changes that occur when annual grasses invade shrub-steppe communities

Changes in the quantity and quality of plant litter occur in many ecosystems as they are invaded by exotic species, which impact soil nutrient cycling and plant community composition. Such changes in sagebrush-steppe communities are occurring with invasion of annual grasses (AG) into a perennial grass (PG) dominated system. We conducted a 5-year litter manipulation study located in the northern Great Basin, USA. Springtime litter was partially or completely removed in three communities with differing levels of invasion (invaded, mixed, and native) to determine how litter removal and litter biomass affected plant-available soil N and plant community composition. Litter biomass (prior to the removal treatment) was negatively correlated with plant-available N in the invaded community, but was positively correlated in the native community. Plant-available N had greater intra- and inter-annual fluctuations in the invaded compared to the mixed or native communities, but was not generally affected by removal treatments. Litter removal had negative effects on AG cover during a warm/dry year and negative effects on PG cover during a cool/wet year in the mixed community. Overall, the effectiveness of springtime litter manipulations on plant-available N were limited and weather dependent, and only removal treatments >75 % had effects on the plant community. Our study demonstrates how communities invaded by AGs have significantly increased temporal variability in nutrient cycling, which may decrease ecosystem stability. Further, we found that the ecological impacts from litter manipulation on sagebrush communities were dependent on the extent of AG invasion, the timing of removal, and seasonal precipitation.     

Mechanisms underlying the impacts of exotic annual grasses in a coastal California meadow

Biological invasions can impact the abundance and diversity of native species, but the specific mechanisms remain poorly discerned. In California grasslands, invasion by European annual grasses has severely reduced the quality of habitat for native forb species. To understand how introduced grasses suppress native and exotic forbs, we examined the response of a Southern California grassland community to factorial removals of live grass and the litter produced in previous seasons. To examine the role that belowground competition for water plays in mediating the impact of grasses, we crossed grass and litter removal treatments with water addition. Our results show that forbs were almost equally suppressed by both competition from live grass and direct interference by litter. Water addition did not ameliorate the effect of grass competition, suggesting that water was not the resource for which plants compete. This evidence is consistent with the susceptibility of forbs to light limitation, especially considering that litter does not consume water or nutrients. Interestingly, despite different histories of co-occurrence with annual grass dominants, native and exotic forbs were comparably suppressed by exotic grasses. Our results indicate that suppression by both live and dead stems underlie the influence of exotic grasses on forb competitors.     

Direct and indirect effects of exotic annual grasses on species composition of a South Australian grassland

Abstract Invasion by Mediterranean annual grasses, such as Avena L. spp. and Bronms L. spp, is one of the major threats to temperate perennial grassland. This study investigated the effects of annual grasses and their litter on the species composition of a grassland near Burra, South Australia. The placement of annual grass litter on soil samples in the glasshouse decreased the establishment or growth of several exotic annual dicots. In the field the addition of annual grass litter slightly decreased the frequency of Danthonia Lam. & DC. tussocks. Furthermore, litter strongly reduced the species richness from 13 species in plots with no litter to nine species in plots with the highest litter level, mainly by decreasing the frequency of common exotic dicots. Native dicot frequency similarly appeared to be decreased by litter addition. In addition to the negative effects of their litter, annual grasses also directly competed with perennial grasses. The magnitude of the competitive effect varied systematically along a slope, suggesting that other factors such as soil properties may control competitive inter actions. The biomass of annual grasses also tended to increase with the addition of their own litter. This combination of positive and negative feedback mechanisms suggests that brief periods favourable for annual grasses, either through management changes or environmental conditions, can lead to persistent changes in the species composition of the system.     

Responses of litter decomposition and nutrient release rate to water and nitrogen addition differed among three plant species dominated in a semi-arid grassland

Background and aims

Precipitation and nitrogen (N) deposition are predicted to increase in northern China. The present paper aimed to better understand how different dominant species in semi-arid grasslands in this region vary in their litter decomposition and nutrient release responses to increases in precipitation and N deposition.

Methods

Above-ground litter of three dominant species (two grasses, Agropyron cristatum and Stipa krylovii, and one forb, Artemisia frigida) was collected from areas without experimental treatments in a semi-arid grassland in Inner Mongolia. Litter decomposition was studied over three years to determine the effects of water and N addition on litter decomposition rate and nutrient dynamics.

Results

Litter mass loss and nutrient release were faster for the forb species than for the two grasses during decomposition. Both water and N addition increased litter mass loss of the grass A. cristatum, while the treatments showed no impacts on that of the forb A. frigida. Supplemental N had time-dependent, positive effects on litter mass loss of the grass S. krylovii. During the three-year decomposition study, the release of N from litter was inhibited by N addition for the three species, and it was promoted by water addition for the two grasses. Across all treatments, N and potassium (K) were released from the litter of all three species, whereas calcium (Ca) was accumulated. Phosphorus (P) and magnesium (Mg) were released from the forb litter but accumulated in the grass litter after three years of decomposition.

Conclusions

Our findings revealed that the litter decomposition response to water and N supplementation differed among dominant plant species in a semi-arid grassland, indicating that changes in dominant plant species induced by projected increases in precipitation and N deposition are likely to affect litter decomposition, nutrient cycling, and further biogeochemical cycles in this grassland. The asynchronous nutrient release of different species’ litter found in the present study highlights the complexity of nutrient replenishment from litter decomposition in the temperate steppe under scenarios of enhancing precipitation and N deposition.

     

Combined effects of nitrogen addition and litter manipulation on nutrient resorption of Leymus chinensis in a semi‐arid grassland of northern China

Plant growth in semi‐arid ecosystems is usually severely limited by soil nutrient availability. Alleviation of these resource stresses by fertiliser application and aboveground litter input may affect plant internal nutrient cycling in such regions. We conducted a 4‐year field experiment to investigate the effects of nitrogen (N) addition (10 g N·m^?2·year^?1) and plant litter manipulation on nutrient resorption of Leymus chinensis, the dominant native grass in a semi‐arid grassland in northern China. Although N addition had no clear effects on N and phosphorus (P) resorption efficiencies in leaves and culms, N fertilisation generally decreased leaf N resorption proficiency by 54%, culm N resorption proficiency by 65%. Moreover, N fertilisation increased leaf P resorption proficiency by 13%, culm P resorption proficiency by 20%. Under ambient or enriched N conditions, litter addition reduced N and P resorption proficiencies in both leaves and culms. The response of P resorption proficiency to litter manipulation was more sensitive than N resorption proficiency: P resorption proficiency in leaves and culms decreased strongly with increasing litter amount under both ambient and enriched N conditions. In contrast, N resorption proficiency was not significantly affected by litter addition, except for leaf N resorption proficiency under ambient N conditions. Furthermore, although litter addition caused a general decrease of leaf and culm nutrient resorption efficiencies under both ambient and enriched N conditions, litter addition effects on nutrient resorption efficiency were much weaker than the effects of litter addition on nutrient resorption proficiency. Taken together, our results show that leaf and non‐leaf organs of L. chinensis respond consistently to altered soil N availability. Our study confirms the strong effects of N addition on plant nutrient resorption processes and the potential role of aboveground litter, the most important natural fertiliser in terrestrial ecosystems, in influencing plant internal nutrient cycling.     

Effects of exotic annual grass litter and local environmental gradients on annual plant community structure

Exotic annual grasses have been introduced into many semi-arid ecosystems worldwide, often to the detriment of native plant communities. The accumulation of litter from these grasses (i.e. residual dry biomass) has been demonstrated to negatively impact native plant communities and promote positive feedbacks to exotic grass persistence. More targeted experiments are needed, however, to determine the relative impact of exotic grass litter on plant community structure across local environmental gradients. We experimentally added exotic grass litter to annual forb-dominated open woodland communities positioned along natural canopy cover gradients in southwest Western Australia. These communities are an important component of this region’s plant biodiversity hotspot and are documented to be under threat from exotic annual grasses. After a one-year treatment period, we measured the effects of exotic grass litter, soil properties, and canopy cover on native and exotic species richness and abundance, as well as common species’ biomass and abundances. Plant community structure was more strongly influenced by soil properties and canopy cover than by grass litter. Total plant abundances per plot, however, were significantly lower in litter addition plots than control plots, a trend driven by native species. Exotic grass litter was also associated with lower abundances of one very common native species: Waitzia acuminata. Our results suggest that exotic grass litter limits the establishment of some native species in this system. Over multiple years, these subtle impacts may contribute substantially to the successful advancement of exotic species into this system, particularly in certain microenvironments.     

氮添加对中亚热带杜鹃灌丛凋落物生产和叶分解的影响

凋落物的生产和分解是生态系统养分循环的重要过程,受到大气氮沉降的深刻影响。但目前相关研究主要集中于森林和草地生态系统,氮沉降对灌丛生态系统凋落物养分归还的影响规律尚不清楚。因此选择亚热带分布广泛的杜鹃灌丛为研究对象,进行了为期两年的模拟氮沉降试验。试验设置4个处理:对照(CK, 0 g m-2 a-1)、低氮(LN, 2 g m-2 a-1)、中氮(MN, 5 g m-2 a-1)和高氮(HN, 10 g m-2 a-1)。结果显示:CK、LN、MN和HN 4种处理下,群落年平均凋落物量分别为(1936.54±358.9)、(2541.89±112.5)、(2342.97±519.8)、(2087.22±391.8) kg/hm²,LN、MN和HN处理样地的凋落量分别比对照样地高出32.68%、21.16%和7.93%;凋落叶、花果、凋落枝和其他组分占总凋落量的比例分别为75.75%、15.09%、7.70%和1.45%,不同浓度氮处理下各组分的凋落量均高于对照样地;凋落物组分表现出明显的季节动态:凋落叶在10—11月份达到峰值,凋落枝在10月份达到峰值,花果凋落物则在5月份凋落量最高,不同氮处理下凋落物的季节动态基本一致;白檀凋落叶分解速率显著高于杜鹃,二者分解95%所需时间分别为5.08—11.11 a和7.69—17.65 a,施氮使白檀凋落叶分解周期比对照样地缩短18.18%—54.28%;凋落叶分解过程中,N元素表现为富集-释放模式,P元素表现为富集模式。研究表明,氮添加能够促进群落中白檀凋落叶分解及N、P元素的释放,说明施氮可以调节凋落叶养分释放模式,对灌丛生态系统的养分循环具有调控作用。     

Effects of elevated CO2 and nitrogen fertilization pretreatments on decomposition on tallgrass prairie leaf litter

Standing dead and green foliage litter was collected in early November 1990 from Andropogon gerardii (C₄), Sorghastrum nutans (C₄), and Poa pratensis (C₃) plants that were grown in large open-top chambers under ambient or twice ambient CO₂ and with or without nitrogen fertilization (45 kg N ha⁻¹). The litter was placed in mesh bags on the soil surface of pristine prairie adjacent to the growth treatment plots and allowed to decay under natural conditions. Litter bags were retrieved at fixed intervals and litter was analyzed for mass loss, carbon chemistry, and total Kjeldahl nitrogen and phosphorus. The results indicate that growth treatments had a relatively minor effect on the initial chemical composition of the litter and its subsequent rate of decay or chemical composition. This suggests that a large indirect effect of CO₂ on surface litter decomposition in the tallgrass prairie would not occur by way of changes in chemistry of leaf litter. However, there was a large difference in characteristics of leaf litter decomposition among the species. Poa leaf litter had a different initial chemistry and decayed more rapidly than C₄ grasses. We conclude that an indirect effect of CO₂ on decomposition and nutrient cycling could occur if CO₂ induces changes in the relative aboveground biomass of the prairie species.     

Response of semi-desert grasslands invaded by non-native grasses to altered disturbance regimes

Aim Using a long‐term data set we investigated the response of semi‐desert grasslands to altered disturbance regimes in conjunction with climate patterns. Specifically, we were interested in the response of a non‐native grass (Eragrostis lehmanniana), mesquite (Prosopis velutina), and native species to the reintroduction of fire and removal of livestock. Location The study site is located on the 45,360‐ha Buenos Aires National Wildlife Refuge (31°32′ N, 110°30′ W) in southern Arizona, USA. In 1985, livestock were removed and prescribed fires were reintroduced to this semi‐desert grassland dominated by non‐native grasses and encroaching mesquite trees. Methods Plant species cover was monitored along 38, 30‐m transects five times over a period of 15 years. Data were analysed using principal components analysis on the variance–covariance and correlation matrix, multivariate analysis of variance for changes over time in relation to environmental data, and analysis of variance for altered disturbance regimes. Results Reintroduction of fire and removal of livestock have not led to an increase in native species diversity or a decrease in non‐native grasses or mesquite. The cover of non‐native grass was influenced by soil type in 1993. Main conclusions Variability of plant community richness, diversity, and cover over time appear to be most closely linked to fluctuations in precipitation rather than human‐altered disturbance regimes. The effects of altered grazing and fire regimes are likely confounded by complex interactions with climatic factors in systems significantly altered from their original physiognomy.     

Litter mixing significantly affects decomposition in the Hulun Buir meadow steppe of Inner Mongolia,China

Aims We explored the decomposition rates of single- and mixed-species litter, the litter-mixing effect and the effect of component litters in a mixture on decomposition.Methods In a litter bag experiment, shoot litters from two dominant grasses (Leymus chinensis and Stipa baicalensis) and one legume (Melissitus ruthenica) were decomposed separately and as a mixture from May 2010 to September 2011 in the Hulun Buir meadow steppe of Inner Mongolia, China. We separated the litter mixture into its individual component litters (i.e. the different single-species litters) and analyzed the changes in litter mass remaining and litter nitrogen (N) remaining during single- and mixed-species litter decomposition.Important findings (i) Litter mixing had significant positive effects on litter decomposition. The litter-mixing effect was strongest for the mixture of S. baicalensis and L. chinensis litters, followed by the mixture of S. baicalensis and M. ruthenica litters. (ii) Single-species component litters decomposed faster in the mixtures than separately (positive effect), but these effects were not significant for legume species M. ruthenica litter. Relative to single-species litter decomposition, the decomposition rates of the two grass (S. baicalensis and L. chinensis) litters significantly increased when they were mixed with each other or with M. ruthenica litter. (iii) For each species litter type, the percentage of litter N remaining during decomposition (N R) differed between the single-species litter and mixed litter treatments. The N R of S. baicalensis litter was higher when it was decomposed in the mixture than in isolation. However, the N R of L. chinensis litter was lowest in its mixture with M. ruthenica among the treatments. Regardless of its decomposition in the mixture or in isolation, the N R of M. ruthenica litter varied little among treatments. There was a significant positive relationship between the N R and percentage of initial litter mass remaining in both the single litter and mixed litter treatments. These results suggest that N transfer may happen among component litters in mixture and further affect the decomposition.     

Soil ecosystem function under native and exotic plant assemblages as alternative states of successional grasslands

Old fields often become dominated by exotic plants establishing persistent community states. Ecosystem functioning may differ widely between such novel communities and the native-dominated counterparts. We evaluated soil ecosystem attributes in native and exotic (synthetic) grass assemblages established on a newly abandoned field, and in remnants of native grassland in the Inland Pampa, Argentina. We asked whether exotic species alter soil functioning through the quality of the litter they shed or by changing the decomposition environment. Litter decomposition of the exotic dominant Festuca arundinacea in exotic assemblages was faster than that of the native dominant Paspalum quadrifarium in native assemblages and remnant grasslands. Decomposition of a standard litter (Triticum aestivum) was also faster in exotic assemblages than in native assemblages and remnant grasslands. In a common garden, F. arundinacea showed higher decay rates than P. quadrifarium, which reflected the higher N content and lower C:N of the exotic grass litter. Soil respiration rates were higher in the exotic than in the native assemblages and remnant grasslands. Yet there were no significant differences in soil N availability or net N mineralization between exotic and native assemblages. Our results suggest that exotic grass dominance affected ecosystem function by producing a more decomposable leaf litter and by increasing soil decomposer activity. These changes might contribute to the extended dominance of fast-growing exotic grasses during old-field succession. Further, increased organic matter turnover under novel, exotic communities could reduce the carbon storage capacity of the system in the long term.     

Effectiveness of repeated autumn and spring fires for understorey restoration in weed‐invaded temperate eucalypt woodlands

Question. Can strategic burning, targeting differing ecological characteristics of native and exotic species, facilitate restoration of native understorey in weed‐invaded temperate grassy eucalypt woodlands? Location. Gippsland Plains, eastern Victoria, Australia. Methods. In a replicated, 5‐year experimental trial, the effects of repeated spring or autumn burning were evaluated for native and exotic plants in a representative, degraded Eucalyptus tereticornis grassy woodland. Treatments aimed to reduce seed banks and modify establishment conditions of exotic annual grasses, and to exhaust vegetative reserves of exotic perennial grasses. Treatments were applied to three grassland patch types, dominated by the native grass Austrodanthonia caespitosa, ubiquitous exotic annuals, or the common exotic perennial grass Paspalum dilatatum. Results. The dominant native grass Austrodanthonia caespitosa and native forbs were resilient to repeated fires, and target exotic annuals and perennials were suppressed differentially by autumn and spring fires. Exotic annuals were also suppressed by drought, reducing the overall treatment effects but indicating important opportunities for restoration. The initially sparse exotic geophyte Romulea rosea increased in cover with fire and the impact of this species on native forbs requires further investigation. There was minimal increase in diversity of subsidiary natives with fire, probably owing to lack of propagules. Conclusions. While fire is often considered to increase ecosystem invasibility, our study showed that strategic use of fire, informed by the relative responses of available native and exotic taxa, is potentially an effective step towards restoration of weed‐invaded temperate eucalypt woodlands.