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.     

Effects of grazing and fertilization on the relationship between species abundance and functional traits in an alpine meadow community on the Tibetan Plateau

Trait‐based approaches can provide a useful tool for linking plant attributes to community structure and ecosystem function. Seed mass and plant height play important roles in the dynamics of plant communities, but few empirical community level studies have tested this, especially in stressful environments. The aim of the present study was to determine if there is a relationship between functional traits (seed mass and plant height) and changes in species relative abundance (SRA) in response to grazing and fertilization. We measured SRA and plant functional traits for 40 common species in a Tibetan Plateau alpine meadow. In the fertilized meadow, seed mass and plant height was significantly positively correlated with the relative abundance of the species. In the grazed meadow, these variables were significantly negatively correlated. Our results demonstrates that plant functional traits can be used to predict the change of SRA in plant community. Grazing promotes the dominance of small‐seeded and short‐stature species, and fertilization facilitates the occurrence of large‐seeded and tall‐stature species.     

Moderate grazing increased alpine meadow soils bacterial abundance and diversity index on the Tibetan Plateau

The response of grassland soil bacterial community characteristics to different grazing intensities is central ecological topics. However, the underlying mechanisms between bacterial abundance, diversity index, and grazing intensity remain unclear. We measured alpine meadow soil bacterial gene richness and diversity index under four grazing intensities using 16S rDNA sequence analysis on the Tibetan Plateau. The results suggest that extreme grazing significantly decreased alpine meadow both bacterial gene abundance and diversity index (p < .05). The lowest operational taxonomic unit numbers were 3,012 ± 447 copies under heavy grazing in the growing season. It was significantly lower than heavy grazing with approximately 3,958 ± 119 copies (p < .05). The Shannon index for medium and high grazing grassland bacterial diversity was slightly higher than for light grazing in the growing season. Furthermore, the lowest index was approximately 9.20 ± 0.50 for extreme grazing of grassland in the growing season. The average bacterial gene abundance and diversity index in the dormancy period were slightly higher than that in the growing season. Soil bulk density, pH, ammonium, and nitrate nitrogen were the main positive factors driving grazed grassland bacterial communities. Our study provides insight into the response of alpine meadows to grazing intensity, demonstrating that moderate grazing increases bacterial community diversity in grazed grasslands.     

The effects of fertilization on the trait-abundance relationships in a Tibetan alpine meadow community

Aims Comparisons of the trait–abundance relationships from various habitat types are critical for community ecology, which can offer us insights about the mechanisms underlying the local community assembly, such as the relative role of neutral vs. niche processes in shaping community structure. Here, we explored the responses of trait–abundance relationships to nitrogen (N), phosphorus (P) and potassium (K) fertilization in an alpine meadow.Methods Five fertilization treatments (an unfertilized control and additions of N, P, K and NPK respectively) were implemented using randomized block design in an alpine Tibetan meadow. Species relative abundance (SRA), plant above-ground biomass and species richness were measured in each plot. For 24 common species, we measured species functional traits: saturated height, specific leaf area (SLA) and leaf dry matter content (LDMC) in each treatment but seed size only in the unfertilized control. Standard major axis (SMA) regression and phylogenetically independent contrasts (PICs) analysis were used to analyse species trait–abundance relationships in response to different fertilization treatments.Important findings Positive correlations between SRA and saturated height were raised following N, P and NPK fertilizations, which indicated an increase in light competition in these plots. In P fertilized plots, SRA was also positively correlated with LDMC because tall grasses with a nutrients conservation strategy often have a relative competitive advantage in capturing limited light and soil nutrients. In K fertilized plots, neither the trait–abundance relationships nor above-ground biomass or species richness significantly differed from that in the control, which suggests that K was not a limiting resource in our study site. These significant correlations between species traits and relative abundance in fertilized treatment suggest that trait-based selection plays an important role in determining species abundance within local communities in alpine meadows.     

高寒草甸物种多样性对藏系牧羊放牧行为的影响

为探究草原植物物种多样性对家畜放牧行为的影响及其机制,在青藏高原高寒草甸开展藏系牧羊轮牧试验,调查植被物种多样性,观察藏系牧羊采食速率、觅食速率和采食时间,并计算藏系牧羊日采食量。结果表明：两年间,植物物种丰富度与藏系牧羊采食速率呈显著正相关关系（P<0.05）;觅食速率、采食时间和日采食量对放牧率响应敏感（P<0.05）,呈夏秋增冬春减的趋势。植物Shannon-Wiener指数与藏系牧羊的采食时间显著负相关（P<0.05）;在暖季或8羊/hm²放牧率下植物Shannon-Wiener指数与藏系牧羊日采食量呈显著正相关（P<0.05）。Pilelou均匀度指数与藏系牧羊采食速率和采食时间显著负相关（P<0.05）;在暖季或8羊/hm²放牧率下Pilelou均匀度指数与藏系牧羊日采食量呈显著负相关（P<0.05）。植物物种丰富度对藏系牧羊放牧行为贡献较大,且放牧藏系牧羊的采食速率和采食时间比觅食速率和日采食量对植物物种丰富度响应更敏感,以用植物物种丰富度为自变量可以更好预测藏系牧羊放牧行为。放牧管理通过影响植被物种多样性从而进一步影响了藏系牧羊放牧行为。放牧行为不仅是评价草地营养价值和家畜生产力的关键指标,也是草地健康管理的基础。因此,明确草原植物物种多样性-藏系牧羊放牧行为的互作机制有助于更好的提高藏系牧羊地生产力,维护草原生态健康。     

施肥和放牧对青藏高原高寒草甸物种丰富度的影响

在草地生态系统中, 施肥通常会导致生物多样性下降, 但是关于引起生物多样性下降的机制还存在着很大的争议。该研究基于一个4年的施肥实验, 试图通过个体大小的不整齐性和单位植物氮含量, 定性地揭示青藏高原东部高寒草甸施肥后多样性下降的原因。研究显示: 在封育地, 施肥致使个体大小不整齐性增加了15%, 并不同程度地增加了物种的高度。同时, 施肥使物种间单位植物氮含量存在显著差异的数目降低了65%。施肥后光竞争加剧, 导致大个体植物排斥小个体植物, 进而引起了物种丰富度下降29.6%。与封育地不同, 放牧地施肥并没有改变个体大小不整齐性和物种的高度, 而是使物种间单位植物氮含量存在显著差异的数目增加了11.4%。施肥并没有改变放牧地的光竞争强度, 而是增加了物种间对土壤营养元素氮的竞争强度, 进而引起了物种丰富度下降17.3%。该研究还发现, 放牧施肥地的物种丰富度下降速度和等级显著低于封育施肥地的物种丰富度下降速度和等级, 这表明放牧减缓了施肥对物种丰富度的影响力。     

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.     

放牧作用下高寒草甸群落物种分布与土壤因子的关系

以青藏高原东北缘高寒草甸为对象,研究不同放牧强度下植物群落和土壤因子的变化,以及群落物种分布与土壤物理结构和化学养分因子的定量关系.结果表明: 放牧导致优势种为垂穗披碱草和大针茅的原植物群落发生分异,高强度放牧样地优势种变为矮生嵩草和阴山扁蓿豆,低强度放牧样地变为垂穗披碱草和冷地早熟禾.随放牧强度增加,物种丰富度、重要值和生物量均显著降低.各放牧强度样地重要值的物种序列均可用对数模型进行拟合;随放牧强度增加,植物重要值累积到占整个群落重要值50%时,需要的物种数降低.土壤速效P、速效K、紧实度、含水量、稳定入渗速率和大团聚体指标随放牧强度显著变化,但变化规律不一致.CCA排序表明,土壤紧实度是放牧作用下影响群落物种分布格局的最关键因子.方差分解表明,土壤因子共解释群落物种分布变异的30.5%,其中土壤物理性状单独解释群落物种分布的22.8%,对群落物种分布的贡献率最高,主要影响放牧干扰下高寒草甸植物群落物种的分布格局.     

Responses of biotic interactions of dominant and subordinate species to decadal warming and simulated rotational grazing in Tibetan alpine meadow

Warming increases competition among plant species in alpine communities by ameliorating harsh environmental conditions, such as low temperatures. Grazing, as the main human activity, may mitigate the effect of warming, as previously reported. However, it is critical to refine the effects of warming on biotic interactions among species, for example, by taking the competitive ability of species into consideration. Based on a 10-year warming and grazing experiment in a Tibetan alpine meadow, we evaluated interspecific biotic interactions of dominant and subordinate species, using the approach of interspecific spatial associations. Warming significantly increased competition between subordinate and dominant species as well as among subordinate species, but not among dominant species. Moreover, facilitation of dominant-subordinate species also increased under warming. Simulated rotational grazing had similar effects to warming, with increasing interspecific competition. Our results show that, when studying the effects of warming on biotic interactions among species, it is necessary to characterize different species pairs relative to their competitive ability, and that simulated rotational grazing does not mitigate the effects of warming in the long term. Our results also provide evidence that the spatial pattern of species is a critical mechanism in species coexistence.     

Different categories of biodiversity explain productivity variation after fertilization in a Tibetan alpine meadow community

The relationship between productivity and biodiversity has long been an important issue in ecological research. However, in recent decades, most ecologists have primarily focused on species diversity while paying little attention to functional diversity and phylogenetic diversity (PD), especially in alpine meadow communities following fertilization. In this study, a fertilization experiment involving the addition of nitrogen, phosphorus, and a mixture of both was implemented in an alpine meadow on the Tibetan Plateau. Species diversity, functional diversity, and PD were measured, and the responses of these parameters to the variation in productivity were analyzed. We found that the productivity of alpine plant communities was colimited by N and P, with N being the principal and P being the secondary limiting nutrient. Our results supported the prediction of both the mass ratio hypothesis and niche complementarity hypothesis in fertilized communities, but these hypotheses were not mutually exclusive. The combination of different aspects of biodiversity not only provides a crucial tool to explain the variation in productivity and to understand the underlying mechanisms but also plays an important role in predicting the variation in productivity of alpine meadow communities, which are sensitive to nutrient enrichment in the context of global change.     

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.     

Methanotrophic community structure and activity under warming and grazing of alpine meadow on the Tibetan Plateau

Knowledge about methanotrophs and their activities is important to understand the microbial mediation of the greenhouse gas CH₄ under climate change and human activities in terrestrial ecosystems. The effects of simulated warming and sheep grazing on methanotrophic abundance, community composition, and activity were studied in an alpine meadow soil on the Tibetan Plateau. There was high abundance of methanotrophs (1.2–3.4 × 10⁸ pmoA gene copies per gram of dry weight soil) assessed by real-time PCR, and warming significantly increased the abundance regardless of grazing. A total of 64 methanotrophic operational taxonomic units (OTUs) were obtained from 1,439 clone sequences, of these OTUs; 63 OTUs (98.4%) belonged to type I methanotrophs, and only one OTU was Methylocystis of type II methanotrophs. The methanotroph community composition and diversity were not apparently affected by the treatments. Warming and grazing significantly enhanced the potential CH₄ oxidation activity. There were significantly negative correlations between methanotrophic abundance and soil moisture and between methanotrophic abundance and NH₄–N content. The study suggests that type I methanotrophs, as the dominance, may play a key role in CH₄ oxidation, and the alpine meadow has great potential to consume more CH₄ under future warmer and grazing conditions on the Tibetan Plateau.     

短期氮素添加和模拟放牧对青藏高原高寒草甸生态系统呼吸的影响

氮沉降和放牧是影响草地碳循环过程的重要环境因子,但很少有研究探讨这些因子交互作用对生态系统呼吸的影响。在西藏高原高寒草甸地区开展了外源氮素添加与刈割模拟放牧实验,测定了其对植物生物量分配、土壤微生物碳氮和生态系统呼吸的影响。结果表明:氮素添加显著促进生态系统呼吸,而模拟放牧对其无显著影响,且降低了氮素添加的刺激作用。氮素添加通过提高微生物氮含量和土壤微生物代谢活性,促进植物地上生产,从而增加生态系统的碳排放;而模拟放牧降低了微生物碳含量,且降低了氮素添加的作用,促进根系的补偿性生长,降低了氮素添加对生态系统碳排放的刺激作用。这表明,放牧压力的存在会抑制氮沉降对高寒草甸生态系统碳排放的促进作用,同时外源氮输入也会缓解放牧压力对高寒草甸生态系统生产的负面影响。     

Linking flowering and reproductive allocation in response to nitrogen addition in an alpine meadow

Aims Plants can change in phenology and biomass allocation in response to environmental change. It has been demonstrated that nitrogen is the most limiting resource for plants in many terrestrial ecosystems. Previous studies have usually focused on either flowering phenology or biomass allocation of plants in response to nitrogen addition; however, attempts to link flowering phenology and biomass allocation are still rare. In this study, we tested the effects of nitrogen addition on both flowering phenology and reproductive allocation in 34 common species. We also examined the potential linkage between flowering time and reproductive allocation in response to nitrogen addition.Methods We conducted a 3-year nitrogen addition experiment in Tibetan alpine meadow. We measured first flowering date and the reproductive allocation for 34 common plant species in control, low and high nitrogen added plots, respectively. One-way analysis of variance was used to examine differences of first flowering date and reproductive allocation among treatments. The relationships between the change in species first flowering date and change in reproductive allocation in response to nitrogen addition were examined by calculating Pearson correlation coefficients.Important findings For most species, both first flowering date and reproductive allocation significantly responded to nitrogen addition. Nitrogen addition significantly delayed the first flowering date and reduced the reproductive allocation for all graminoid species, but accelerated flowering and increased reproductive allocation for most forb species. We found that changes in first flowering date significantly negatively correlated with the changes in reproductive allocation over species in response to nitrogen, which indicated a positive relationship between flowering response and plant performance in reproductive allocation. Species that advanced their flowering time with nitrogen addition increased their reproductive allocation, whereas those that delayed flowering time tended to decline in reproductive allocation with nitrogen addition. Our results suggest that species-specific switch from vegetative growth to reproductive growth could influence species performance.     

Relationship between species relative abundance and plant traits for an infertile habitat

This study tested whether differences in species abundance at an infertile site could be explained by differences in the species' plant traits. Nine traits were chosen for the analysis based on results of previous studies conducted across soil fertility gradients. The traits were measured for each of seven herbaceous species whose abundance ranged from 5% to 100% of locations occupied in a ridgetop habitat. Using linear regression, significant relationships were found between species relative abundance and each of five traits. In these relationships, a trait explained between 69% and 88% of interspecific variation in abundance. Relatively abundant species had a slower growth rate, smaller shoot mass, higher root to shoot ratio, slower loss of leaf tissue to herbivores and higher infection of roots by mycorrhizal fungi than less abundant species. Using three of these five traits (i.e. shoot mass, mycorrhizal infection and loss of leaf tissue to herbivores) as independent variables in a multiple regression equation explained 99% of interspecific variation in abundance. The latter result indicates that species relative abundance can be explained for a single habitat by choosing traits found to be related to species abundance in previous gradient studies. However, not every trait chosen was significantly related to species abundance. Therefore, a large number of traits may have to be chosen initially to ensure that some subset of these traits can explain species relative abundance.     

放牧对青藏高原东缘高寒草甸群落27种植物地上生物量分配的影响

通过对比研究青藏高原高寒草甸27种植物群落组分种在放牧和长期排除放牧生境中的生物量分配差异,试图揭示长期放牧干扰对植物生活史对策的影响.结果表明:(1)放牧对群落物种个体生物量大小和生物量分配有着显著的影响;(2)总体来看:多数物种(24)放牧生境中的平均个体生物量明显小于禁牧地中的平均个体生物量;而多数物种在放牧后(23种)繁殖分配明显增加;茎分配有增有减(15减小12种增加);叶分配呈减小趋势(20种减小7增加).(3)放牧的影响在不同物种间和功能群间都存在着明显的差异.放牧使毒草茎分配减小叶分配增加,繁殖分配几乎不受影响;豆科和杂草繁殖分配增加,茎分配和叶分配减小,其中豆科两个种的生物量分配变化都不显著;禾草叶分配减小,繁殖分配和茎分配增加; (4)在群落水平上,放牧使繁殖分配和叶分配增加,茎分配减少.     

施肥和刈割对青藏高原东部高寒草甸群落生物量补偿效应的影响

通过对青藏高原东部高寒草甸植物群落在4个施肥处理及7个刈割水平下的群落生物量、物种丰富度和补偿指数的比较,分析了该类草甸群落主要特征对施肥及刈割强度差异的响应.结果表明:施肥显著增加了群落生物量,并降低物种丰富度,而刈割处理虽然增加了物种丰富度,但对群落生物量无显著影响;随着施肥量的增加,高寒草甸由莎草科和杂草类物种占优势的群落构成逐渐向禾本科物种占优势的群落构成转变;施肥可提高群落的补偿能力,在不同施肥梯度中,刈割后植物群落补偿能力也不相同;随着施肥量的增加,增加刈割强度,减少刈割次数,发生超补偿的机率增大;可见,施肥是主效应,刈割对群落主要特征的影响随营养水平的不同而发生变化.     

放牧干扰下高寒草甸物种、生活型和功能群多样性与生物量的关系

为明晰放牧强度对高寒草甸物种、生活型以及功能群多样性和生物量的时间累积作用以及对多样性与生物量之间相互关系的影响。选择青藏高原东北缘高寒草甸,在6个放牧强度样地连续4 a(2012—2015)进行物种、生活型、功能群多样性和生物量调查。采用重复测量方差分析和线性回归法分析放牧强度和放牧年份对物种、生活型、功能群多样性以及多样性与生物量之间的相关关系的影响。结果表明:(1)放牧强度和放牧年份均对生物量产生显著影响。随放牧强度增加,生物量显著降低。但随放牧年份延长,不同放牧强度区生物量无一致的变化规律。(2)放牧年份对物种、生活型和功能群丰富度、均匀度和优势度的影响均达到显著水平。但放牧强度和放牧年份只对物种丰富度产生交互作用,放牧强度对物种丰富度的影响具有时间的累积效应。(3)放牧干扰下不同层次多样性,仅物种丰富度与所有多样性指数显著相关,物种丰富度可以作为物种多样性测度的代表性指标。(4)多样性与生物量关系的研究,对丰富度而言,仅低放牧强度样地(Plot2)中物种丰富度随生物量增加而显著降低,其余放牧强度样地的物种、生活型和功能群丰富度均与生物量无关。对均匀度而言,高放牧强度样地(Plot4、Plot5、Plot6)生活型均匀度随生物量的增加而显著降低。对优势度而言,高放牧强度样地生活型优势度随生物量的增加而显著增加。生活型多样性可作为放牧干扰下生物量变化快速预测的有效指标。     

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。