Litter species traits, but not richness, contribute to carbon and nitrogen dynamics in an alpine meadow on the Tibetan Plateau

Aims

Litter, as afterlife of plants, plays an important role in driving belowground decomposition processes. Here we tested effects of litter species identity and diversity on carbon (C) and nitrogen (N) dynamics during litter decomposition in N-limited alpine meadow soil from the Qinghai–Tibet Plateau.

Methods

We incubated litters of four meadow species, a sedge (“S”, Kobresia humilis), a grass (“G”, Elymus nutans), a herb (“H”, Saussurea superba), and a legume (“L”, Oxytropis falcata), in monoculture and in mixture with meadow soil. CO₂ release was measured 21 times during the incubation, and soil available N and microbial biomass C and N were measured before and after the experiment.

Results

The organic C decay rate did not differ much among soils amended with monocultures or mixtures of litter, except in the H, S, L, and S+H treatments, which had much higher decay rates. Potential decomposable C pools were lowest in the control, highest in the L treatment, and intermediate in the S treatment. Mineralized N was completely immobilized by soil microbes in all treatments except the control, S+L, and S+G+L treatments. Litter mixtures had both additive and non-additive effects on CO₂-C emission (mainly antagonistic effects), net N mineralization (mainly synergistic), and microbial biomass C and N (both). Overall, these parameters were not significantly correlated with litter species richness. Similarly, microbial C or N was not significantly correlated with litter N content or C/N. However, cumulative CO₂-C emission and net N mineralization were positively correlated with litter N content and negatively correlated with litter C/N.

Conclusions

Litter N content and C/N rather than litter species richness drove the release of CO₂-C and net available N in this ecosystem. The antagonistic effects of litter mixtures contributed to a modest release of CO₂-C, but their synergistic effects enhanced net available N. We suggest that in alpine meadow communities, balancing species with high and low N contents will benefit soil carbon sequestration and plant competition for available N with soil microbes.     

Grazing alters species relative abundance by affecting plant functional traits in a Tibetan subalpine meadow

Domestic livestock grazing has caused dramatic changes in plant community composition across the globe. However, the response of plant species abundance in communities subject to grazing has not often been investigated through a functional lens, especially for belowground traits. Grazing directly impacts aboveground plant tissues, but the relationships between above‐ and belowground traits, and their influence on species abundance are also not well known. We collected plant trait and species relative abundance data in the grazed and nongrazed meadow plant communities in a species‐rich subalpine ecosystem of the Qinghai–Tibet Plateau. We measured three aboveground traits (leaf photosynthesis rate, specific leaf area, and maximum height) and five belowground traits (root average diameter, root biomass, specific root length, root tissue density, and specific root area). We tested for shifts in the relationship between species relative abundance and among all measured traits under grazing compared with the nongrazed meadow. We also compared the power of above‐ and belowground traits to predict species relative abundance. We observed a significant shift from a resource conservation strategy to a resource acquisition strategy. Moreover, this resource conservation versus resource acquisition trade‐off can also determine species relative abundance in the grazed and nongrazed plant communities. Specifically, abundant species in the nongrazed meadow had aboveground and belowground traits that are associated with high resource conservation, whereas aboveground and belowground traits that are correlated with high resource acquisition determined species relative abundance in the grazed meadow. However, belowground traits were found to explain more variances in species relative abundance than aboveground traits in the nongrazed meadow, while aboveground and belowground traits had comparable predictive power in the grazed meadow. We show that species relative abundance in both the grazed and the nongrazed meadows can be predicted by both aboveground traits and belowground traits associated with a resource acquisition versus conservation trade‐off. More importantly, we show that belowground traits have higher predictive power of species relative abundance than aboveground traits in the nongrazed meadow, whereas in the grazed meadows, above‐ and belowground traits had comparable high predictive power.     

Soil warming increases plant species richness but decreases germination from the alpine soil seed bank

Global warming is occurring more rapidly above the treeline than at lower elevations and alpine areas are predicted to experience above average warming in the future. Temperature is a primary factor in stimulating seed germination and regulating changes in seed dormancy status. Thus, plant regeneration from seed will be crucial to the persistence, migration and post disturbance recruitment of alpine plants in future climates. Here, we present the first assessment of the impact of soil warming on germination from the persistent alpine soil seed bank. Contrary to expectations, soil warming lead to reduced overall germination from the soil seed bank. However, germination response to soil temperature was species specific such that total species richness actually increased by nine with soil warming. We further explored the system by assessing the prevalence of seed dormancy and germination response to soil disturbance, the frequency of which is predicted to increase under climate change. Seeds of a significant proportion of species demonstrated physiological dormancy mechanisms and germination of several species appeared to be intrinsically linked to soil disturbance. In addition, we found no evidence of subalpine species and little evidence of exotic weed species in the soil, suggesting that the soil seed bank will not facilitate their invasion of the alpine zone. In conclusion, changes in recruitment via the alpine soil seed bank can be expected under climate change, as a result of altered dormancy alleviation and germination cues. Furthermore, the alpine soil seed bank, and the species richness therein, has the potential to help maintain local species diversity, support species range shift and moderate species dominance. Implications for alpine management and areas for further study are also discussed.     

高寒草甸植物群落功能多样性对不同生长期干旱的响应机制

植物对干旱胁迫的响应表现为各功能性状的差异化表达。全球气候变化下,青藏高原地区降水格局发生改变,高寒草甸群落功能性状及功能多样性在不同生长期干旱事件下的响应机制对加深高寒草甸适应气候变化认知具有重要意义。以藏北高寒草甸为研究对象,设置截雨棚模拟生长季前期（ED）、中期（MD）和非生长季时期（ND）干旱事件,通过观测群落物种功能性状,分析高寒草甸群落功能多样性对不同生长期干旱的响应机制。结果表明：（1）叶片功能性状对干旱存在差异响应,表现为叶片小而厚且寿命长,同化速率降低,并受氮元素限制加剧;（2）生长季前期干旱对高寒草甸群落功能性状的影响最强,生长季中期干旱次之,非生长季干旱的影响最弱;（3）生长季干旱处理显著改变了群落的功能多样性,ED处理下功能分散度指数（FDiv）和功能分异度指数（FDis）显著降低（P<0.05）,而Rao二次熵指数（RaoQ）显著升高（P<0.05）,MD处理下功能均匀度指数（FEve）显著降低（P<0.05）;（4）相关性分析得出,群落功能性状与功能多样性对干旱的响应之间存在着联系。本研究发现高寒草甸植物功能性状与群落功能多样性对生长季前期和中期干旱存在差异化响应,指示着高寒草甸植物群落在响应不同时期干旱时可能采取不同的生存策略,即对生长季前期干旱采用耐旱策略、对生长季中期干旱采用避旱策略。探讨了高寒草甸植物群落功能多样性对不同生长时期干旱胁迫的响应机制,为预测未来季节性干旱事件对青藏高原高寒草甸植物功能性状、群落特征和功能多样性的影响提供科学依据。     

藏北高寒草甸植物群落对土壤线虫群落功能结构的影响

2011年5-11月,对西藏北部高寒草甸3种典型植物群落下0-30cm范围内不同深度土层的土壤线虫群落进行调查,采用浅盆法分离线虫,土壤性质指标,如pH、含水量、电导率分别采用电位法、烘干法、电导率仪法进行测定,应用营养类群组成、c-p类群结构及营养结构特征指数,以及营养类群、c-p类群与土壤性质之间的关系等特征值分析高寒环境下土壤线虫群落的功能结构特征,以了解高寒环境下植被对土壤线虫群落功能结构的影响.调查共分离得到33038条土壤线虫,隶属于2纲6目51科93属;线虫个体密度平均为847条/100g干土;表聚性明显.研究结果表明,高寒草甸不同植物群落的土壤线虫群落营养类群组成及分布特征均存在一定差异,植食性线虫和食细菌性线虫是调查区域的主要营养类群,不同植物群落间植食性线虫和杂食/捕食性线虫的相对多度差异明显.c-p类群组成结构特征结果表明:3种植物群落的土壤线虫cp2类群均为优势类群,生活策略以r-对策为主;高山嵩草植物群落土壤中的线虫食物资源在3种植物群落中最丰富;藏北嵩草群落土壤线虫数量低的可能原因是线虫食物资源的减少限制了cp1、cp2类群的增殖.PPI值表明:委陵菜群落受扰动的影响程度大于其余两种植被类型,而MI、PPI/MI值及cp5类群数量的结果则表明:委陵菜群落的稳定性较高,受到的干扰在3种植物群落中最少.F/B及NCR值均说明了3种植物群落的土壤有机质分解均主要依靠细菌分解途径.相关性分析结果表明:杂食/捕食性线虫在枯草期明显受到土壤含水量的影响;食真菌性线虫与土壤pH之间的关系密切,在盛长期则明显受到土壤电导率的影响;食细菌性线虫仅在返青期与pH有相关性.不同植物群落下土壤线虫群落功能结构特征的分异显示出线虫指示环境因子影响土壤生态系统的潜力.     

高寒草甸不同坡向植物群落物种多样性与功能多样性的关系

植物群落物种多样性与功能多样性是生态学研究的热点问题之一,研究其对于保护生物多样性和维持生态系统功能具有重要意义。采用野外群落调查方法,探讨了物种多样性与功能多样性在坡向梯度上的变化格局及其关联性。结果显示：（1）从北坡到南坡,土壤含水量、有机碳呈递减趋势;坡度、照度、土温、pH呈递增趋势;土壤全氮、全磷在西坡明显低于其他各坡向;（2）物种多样性指数均由北坡向南坡依次递减;（3）一元性状功能多样性在各坡向间的差异显著（P<0.05）,除LDMC_FDrao外,其他功能性状均为北坡大于南坡;（4）多元性状功能丰富度与功能均匀度均由北坡向南坡呈递减的趋势,各坡向的功能均匀度差异性不显著。通过相关分析与通径分析发现,功能多样性指数的主要环境决定因子与限定因子均存在差异：功能丰富度、功能均匀度主要环境限定因子为pH,功能离散度主要环境限定因子为土温;功能离散度主要环境决定因子为土壤含水量;（5）物种多样性指数与功能丰富度呈正线性相关,与功能离散度呈负线性相关,而与功能均匀度不相关。     

Grazing alters environmental control mechanisms of evapotranspiration in an alpine meadow of the Tibetan Plateau

Aims Evapotranspiration (ET) is an important component of the terrestrial water cycle and is easily affected by external disturbances, such as climate change and grazing. Identifying ET responses to grazing is instructive for determining grazing activity and informative for understanding the water cycle.     

Grazing intensifies degradation of a Tibetan Plateau alpine meadow through plant–pest interaction

Understanding the plant–pest interaction under warming with grazing conditions is critical to predict the response of alpine meadow to future climate change. We investigated the effects of experimental warming and grazing on the interaction between plants and the grassland caterpillar Gynaephora menyuanensis in an alpine meadow on the Tibetan Plateau in 2010 and 2011. Our results showed that grazing significantly increased nitrogen concentration in graminoids and sward openness with a lower sward height, sward coverage, and plant litter mass in the community. Grazing significantly increased G. menyuanensis body size and potential fecundity in 2010. The increases in female body size were about twofold greater than in males. In addition, grazing significantly increased G. menyuanensis density and its negative effects on aboveground biomass and graminoid coverage in 2011. We found that G. menyuanensis body size was significantly positively correlated with nitrogen concentration in graminoids but negatively correlated with plant litter mass. Even though warming did not significantly increased G. menyuanensis performance and the negative effects of G. menyuanensis on alpine meadow, the increases in G. menyuanensis growth rate and its negative effect on aboveground biomass under the warming with grazing treatment were significantly higher than those under the no warming with grazing treatment. The positive effects of grazing on G. menyuanensis performance and its damage were exacerbated by the warming treatment. Our results suggest that the fitness of G. menyuanensis would increase under future warming with grazing conditions, thereby posing a greater risk to alpine meadow and livestock production.     

Linking grazing response of species abundance to functional traits in the Tibetan alpine meadow

Whether grazing response of species abundance can be predicted by plant functional trait remains a challenge untested in a specific ecosystem. We measured species abundance and relevant functional traits for 30 common component species in a moderately grazed and a control community over 3 years (2005, 2006 and 2007) in a Kobresia capillifolia dominated alpine meadow on the Tibetan Plateau. Our objective was to examine species response to grazing and to test the relationship between the response of species abundance and functional traits in grazed and control habitats. Our results showed: i) in terms of species relative abundance and saturated height (the maximum height of an adult individual), most component species significantly decrease in response to moderate grazing and the effect differed among species and between functional groups. ii) The response of species abundance was significantly negatively correlated with saturated height, but not correlated with specific leaf area or seed size. We concluded that the response direction of species abundance to grazing can be predicted by functional traits, whereas it is a weak predictor of the extent of grazing response in species abundance.     

高寒草甸不同草地群落物种多样性与生产力关系研究

生态系统的结构和功能、生物多样性与生产力的关系问题是近年来群落生态学中研究的中心问题,其中,生态系统生产力水平是其功能的重要表现形式,用4种不同草地类型探讨自然群落的物种多样性与生产力关系.结果表明,矮嵩草草甸、小嵩草草甸和金露梅灌丛群落中物种多样性与生产力的关系呈线性增加关系,藏嵩草沼泽化草甸群落中线性增加关系不显著,这表明群落生产力除受物种多样性的影响外,也受物种本身特征和环境资源的影响.不同的环境资源和环境异质性是形成群落结构特征、物种多样性分布格局差异的主要原因之一.     

Livestock settlement affects shrub abundance via plant functional diversity but not species richness in arid environments

Plant Ecology - Recent theoretical and empirical studies have assumed that livestock grazing acts as a biotic filter favouring plant species with functional traits that confer resistance to...     

草甸和沼泽植物群落功能多样性与生产力的关系

功能多样性-生产力关系研究结果支持质量比假说和多样性假说, 但对于这两种假说的适用条件尚有争议。通过对吉林省西部草甸和沼泽植物群落的地上生物量、2个物种多样性指标(物种丰富度和Shannon-Weaver指数)、7种植物性状的两类功能多样性指标(群落权重均值和Rao二次熵), 以及土壤环境因子进行调查测量, 研究了群落功能多样性与生产力的关系。结果表明: 1)功能多样性与生产力的关系比物种多样性与生产力的关系更为密切; 2)功能群落权重均值解释生产力变异的能力好于Rao二次熵, 即优势物种对群落生产力的影响作用更大; 3)水淹条件影响着功能多样性与生产力的关系, 以群落权重均值为基础的质量比假说适于解释草甸群落功能多样性与生产力的关系, 而以Rao二次熵为基础的多样性假说适于解释有强烈环境筛(水淹)的沼泽群落功能多样性与生产力的关系。     

Species richness increases the resilience of wetland plant communities in a tropical floodplain

In the last two decades, the relationship between diversity and stability/ecosystem functioning has been widely discussed and has become a central issue in ecology. Here, we assessed the relationship between wetland plant diversity and community resilience after a disturbance. Our study area was located in the Upper Paraná River floodplain (Brazil). An experiment was carried out in situ (18 1 m × 1 m plots with richness varying from 1 to 18 species). In each plot, we recorded the number of species, total per cent vegetation cover and per cent age cover of each species. The above‐ground biomass of wetland plants was removed, simulating a disturbance by animal trampling or an extreme flood. The recovery of vegetation was monitored over 3 months. According to a linear regression, the recovery of wetland plants was positively correlated with diversity. Comparisons with plots containing monocultures of one of the dominant species (Polygonum stelligerum) suggested that this species did not overyield in mixed cultures. Thus, our experiments indicate that the higher resilience in richer plots after a disturbance is mainly due to the fact that species have different resource use requirements (complementarity effect) and not due to the presence of a single, more productive species. Our experiment carried out in a more real condition (in situ) showed that biodiversity is important to wetland functioning and stability, paralleling the results obtained in laboratory and mesocosms experiments. These results also suggest that the loss of plant diversity in our study area could compromise community recovery following strong disturbances.     

高寒草甸不同草地类型功能群多样性及组成对植物群落生产力的影响

对不同类型草地功能群多样性和组成与植物群落生产力之间的关系进行了探讨。结果表明：(1)在矮嵩草(Kobresia humlis)草甸和金露梅(Potentilla froticosa)灌丛中,豆科植物的作用比较明显,而其他功能群植物的作用较弱。(2)在藏嵩草(Kobresia tibetica)沼泽化草甸和小嵩草(K．pygmaca)草甸中,虽然杂类草、C3植物和莎草科植物功能群的生产力占群落初级生产力的比例较大,但二者在统计上没有显著性差异,这表明群落生产力除受物种多样性的影响外,也受物种本身特征和环境资源的影响,更主要的是受到功能群内物种密度和均匀度的影响,即功能群组成比功能群多样性更能说明对生态系统过程的影响。(3)不同类型草地群落植物功能群盖度与群落初级生产力呈显著的线性相关。(4)不同类型草地群落生产力与功能群内物种数的变化均表现为单峰曲线关系,即功能群内物种数处于中间水平时,群落生产力最高。     

Relationship between functional diversity and productivity in meadow and marsh plant communities

功能多样性-生产力关系研究结果支持质量比假说和多样性假说, 但对于这两种假说的适用条件尚有争议。通过对吉林省西部草甸和沼泽植物群落的地上生物量、2个物种多样性指标(物种丰富度和Shannon-Weaver指数)、7种植物性状的两类功能多样性指标(群落权重均值和Rao二次熵), 以及土壤环境因子进行调查测量, 研究了群落功能多样性与生产力的关系。结果表明: 1)功能多样性与生产力的关系比物种多样性与生产力的关系更为密切; 2)功能群落权重均值解释生产力变异的能力好于Rao二次熵, 即优势物种对群落生产力的影响作用更大; 3)水淹条件影响着功能多样性与生产力的关系, 以群落权重均值为基础的质量比假说适于解释草甸群落功能多样性与生产力的关系, 而以Rao二次熵为基础的多样性假说适于解释有强烈环境筛(水淹)的沼泽群落功能多样性与生产力的关系。