Grazing increases functional richness but not functional divergence in Tibetan alpine meadow plant communities

Plant community diversity and ecosystem function are conditioned by competition among co-occurring species for multiple resources. Previous studies suggest that removal of standing biomass by grazing decreases competition for light, but coincident grazing effects on competition for soil nutrients remain largely unknown in Tibetan rangelands where grazing tends to deplete soil phosphorus availability. We measured five functional traits indicative of plant productivity and stoichiometry leaf carbon concentration (LCC), leaf nitrogen concentration (LNC), leaf phosphorus concentration (LPC), specific leaf area (SLA), leaf dry matter content (LDMC) for component species of plant communities in grazed and ungrazed plots in five Tibetan alpine meadows. We examined the diversity of traits singly Rao index of functional diversity (FDrao) and in aggregate functional richness (FRic), functional divergence (FDiv), and functional evenness (FEve) in response to grazing. We tested whether foliar trait diversity increases with nutrient competition but decreases with light competition when competitive exclusion is reduced by grazing. The FDrao of LPC significantly increased under grazing, but FDrao for LCC, LNC and SLA tended to decrease. The FDrao of LDMC increased at the drier site but decreased at the wettest site. There was a strong negative association between increase in FDrao of LPC and decrease in soil nutrients, especially soil phosphorus availability. The FRic for all five traits together increased with species diversity following grazing, but neither FDiv nor FEve differed significantly between grazed and ungrazed plots at most sites. Grazing in Tibetan alpine meadows tends to increase competition for soil phosphorus while decreasing competition for light, resulting in an increase in the functional richness in grazed plant communities without any significant changes in the overall functional diversity of foliar traits. Our study highlights the potential importance of grazing mediated competition for multiple resources in alpine meadow ecosystems.     

Small-scale species richness and its spatial variation in an alpine meadow on the Qinghai-Tibet Plateau

We investigated how the high small-scale species richness of an alpine meadow on the Qinghai-Tibet Plateau, China, is maintained. This area is characterized by strong wind and severe cold during long winters. In winter, most livestock is grazed on dead leaves in small pastures near farmers’ residences, whereas in the short summer, livestock is grazed in mountainous areas far from farmers’ residences. The number of plant species and the aboveground biomass were surveyed for three adjacent pastures differing in grazing management: a late-winter grazing pasture grazed moderately from 1 February to 30 April, an early-winter grazing pasture grazed lightly from 20 September to late October, and a whole-year grazing pasture grazed intensively throughout the entire year. In each pasture, we harvested the aboveground biomass from 80 or 100 quadrats of 0.01 m² along a transect and classified the contents by species. We observed 15.5–19.7 species per 0.01 m², which is high richness per 0.01 m² on a worldwide scale. The species richness in the two winter grazing pastures was higher than that in the whole-year grazing pasture. The spatial variation in species richness and species composition in the two winter grazing pastures in which species richness was high was greater than that in the whole-year grazing pasture in which species richness was lower. Most of the leaves that are preserved on the winter grazing pastures during summer are blown away by strong winds during winter, and the remaining leaves are completely exhausted in winter by livestock grazing. A pasture with a high richess is accompanied by a high spatial variation in species richness and species composition. There is a high possibility that the characteristic of spatial variation is also caused by traditional grazing practices in this area.     

氮素添加对青藏高原高寒草甸植物群落物种丰富度及其与地上生产力关系的影响

植物种群对有限资源的竞争是决定植物群落物种组成、多样性和生产力等群落结构和功能的主要因素。该文以青藏高原高寒草甸为研究对象, 研究了短期内不同水平的氮素添加对高寒草甸植物群落的影响。结果表明: 1)氮素添加提高了土壤中NO₃^--N等可利用资源的含量, 增加了植物群落植被的盖度, 减小了植被的透光率, 随着施氮量的增加, 群落中物种丰富度显著降低(p < 0.001); 2)氮素添加显著改变了植物群落的地上生产力(p < 0.05), 随着施氮量的增加, 地上生产力呈先增加后降低的变化趋势, 各功能群中禾草生物量显著增加, 而杂类草和豆科植物生物量随施氮量的增加逐渐减少; 3)物种多样性与植被透光率呈线性正相关(p < 0.05); 地上生产力与土壤NO₃^--N含量呈线性正相关(p < 0.05); 物种丰富度与地上生产力之间呈负相关关系。这说明短期内氮素添加通过改变土壤中NO₃^--N等可利用资源的含量而对植物群落物种组成和地上生产力产生影响。     

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。     

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.     

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.     

模拟增温对西藏高原高寒草甸土壤供氮潜力的影响

过去几十年青藏高原呈现显著的增温趋势,冬季增温幅度显著高于生长季的季节非对称特征。气候变暖会对生态系统氮素循环产生重要影响,但关于全年增温与冬季增温对高寒生态系统氮循环的不同影响仍缺乏研究。在青藏高原高寒草甸区开展模拟增温试验,研究季节非对称增温对高寒草甸生态系统氮循环的影响。该试验布设于2010年7月,设置3种处理(不增温、冬季增温与全年增温)。研究结果发现,开顶箱增温装置造成了小环境的暖干化:显著提高了地表空气温度和表层土壤温度,降低了表层土壤含水量。冬季增温会加剧土壤中氮素的流失,所以在经历了冬季增温后土壤氮含量显著降低;在生长季节,土壤氮素周转速率受土壤水分的调控,在降雨较少的季节,增温引起的土壤含水量降低会抑制土壤氮周转速率。对于土壤微生物量而言,高寒草甸土壤微生物量碳表现出明显的季节动态,在生长季旺盛期较低,在生长季末期和初冬季节反而较高,这说明为了降低对土壤养分的竞争,高寒草甸植物氮吸收与土壤微生物氮固持在时间上存在分离。研究结果表明,冬季增温导致的土壤养分含量变化会影响随后生长季植物群落的生产力、结构组成与碳氮循环等过程,对生态系统过程产生深远的影响。     

青藏高原高寒草甸生态系统碳增汇潜力

为了揭示青藏高原高寒草甸生态系统植被变化对碳储量的影响,以原生矮嵩草草甸、退化草甸、人工草地以及农田为研究对象,对比分析了该4种不同土地格局下生态系统的有机碳现状.以原生矮嵩草草甸土壤碳储量为基准对不同类型高寒生态系统的碳增汇潜力进行了估算.结果表明:不同类型生态系统的碳储量和碳增汇潜力有很大差异,在0-40cm土层中,(1)原生草甸碳储量最高,达到17098 g C/m2,退化草甸、人工草地和农田的有机碳汇增加潜力分别为:5637、3823、1567 g C/m2.(2)对于退化草甸和人工草地,土壤有机碳含量和密度明显低于原生草甸和农田.(3)地下生物量碳储量主要集中在0-20cm,且原生草甸地下生物量的碳储量比其他3个植被类型高3.6-5倍.总体上,青藏高原草地生态系统存在巨大的碳增汇潜力.     

青藏高原高寒草地植物物种丰富度及其与环境因子和生物量的关系

在青藏高原进行了大范围的群落调查 ,研究高原的两种主要草地群落类型———高寒草甸和高寒草原的植物物种丰富度及其变化。结果表明 :(1)在 5 0个样地 2 5 0个 1m× 1m的样方中 ,共出现 2 6 7种植物 ,其中高寒草甸179种 ,高寒草原 135种。在高寒草甸 ,1m2 样方内物种数最多为 32种 ,最少的仅为 3种 ;在高寒草原 ,物种数最多为 18种 /m2 ,最少的仅为 2种 /m2 。 (2 )物种丰富度随经度和纬度的增加呈增加趋势 ;随海拔的上升呈减少趋势。对物种丰富度与环境因子之间进行逐步回归 ,发现物种丰富度与生长季降水和温暖指数呈显著正相关。 (3)物种丰富度与地上生物量呈显著正相关。     

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。     

Temperature‐mediated responses of carbon fluxes to precipitation variabilities in an alpine meadow ecosystem on the Tibetan Plateau

Effects of climate warming and changing precipitation on ecosystem carbon fluxes have been intensively studied. However, how they co‐regulate carbon fluxes is still elusive for some understudied ecosystems. To fill the gap, we examined net ecosystem productivity (NEP), gross ecosystem productivity (GEP,) and ecosystem respiration (ER) responses to multilevel of temperature increments (control, warming 1, warming 2, warming 3, warming 4) in three contrasting hydrological growing seasons in a typical semiarid alpine meadow. We found that carbon fluxes responded to precipitation variations more strongly in low‐level warming treatments than in high‐level ones. The distinct responses were attributable to different soil water conditions and community composition under low‐level and high‐level warming during the three growing seasons. In addition, carbon fluxes were much more sensitive to decreased than to increased precipitation in low‐level warming treatments, but not in high‐level ones. At a regional scale, this negative asymmetry was further corroborated. This study reveals that future precipitation changes, particularly decreased precipitation would induce significant change in carbon fluxes, and the effect magnitude is regulated by climate warming size.     

若尔盖草甸退化对土壤碳、氮和碳稳定同位素的影响

为了解不同退化阶段高寒草甸土壤碳、氮和碳稳定同位素的差异,对若尔盖湿地内沼泽草甸、草原化草甸、退化草甸3个阶段土壤的碳、氮和碳稳定同位素进行了分析.结果表明:若尔盖湿地草甸土壤δ¹³C 值介于-26.21‰～-24.72‰之间,土壤δ¹³C 值随土层加深而增大.土壤δ¹³C 值与有机碳含量对数值呈线性负相关.表层土壤(0～10 cm)δ¹³C值大小顺序为草原化草甸>退化草甸>沼泽草甸,β值大小顺序为草原化草甸>沼泽草甸>退化草甸.沼泽草甸、草原化草甸、退化草甸0～30 cm 土壤碳含量分别为105.32、42.11和31.12 g·kg^-1,氮含量分别为8.74、3.41和2.81 g·kg^-1,C/N分别为11.26、11.23和10.89.随着草甸的退化,土壤碳、氮呈降低趋势,退化草甸C/N值低于沼泽草甸和草原化草甸.随着土层深度加深,碳、氮含量呈现降低趋势.草甸退化导致的土壤δ¹³C 值差异主要发生在表层0～10 cm.3个退化阶段中,退化草甸土壤的β值和C/N最低,表明退化草甸土壤矿化作用较强.     

Effects of resource additions on species richness and ANPP in an alpine meadow community

Aims Theories based on resource additions indicate that plant species richness is mainly determined by the number of limiting resources. However, the individual effects of various limiting resources on species richness and aboveground net primary productivity (ANPP) are less well understood. Here, we analyzed potential linkages between additions of limiting resources, species loss and ANPP increase and further explored the underlying mechanisms.Methods Resources (N, P, K and water) were added in a completely randomized block design to alpine meadow plots in the Qinghai-Tibetan Plateau. Plant aboveground biomass, species composition, mean plant height and light availability were measured in each plot. Regression and analysis of variance were used to analyze the responses of these measures to the different resource-addition treatments.Important findings Species richness decreased with increasing number of added limiting resources, suggesting that plant diversity was apparently determined by the number of limiting resources. Nitrogen was the most important limiting resource affecting species richness, whereas P and K alone had negligible effects. The largest reduction in species richness occurred when all three elements were added in combination. Water played a different role compared with the other limiting resources. Species richness increased when water was added to the treatments with N and P or with N, P and K. The decreases in species richness after resource additions were paralleled by increases in ANPP and decreases in light penetration into the plant canopy, suggesting that increased light competition was responsible for the negative effects of resource additions on plant species richness.     

青藏高原高寒草甸生态系统净二氧化碳交换量特征

高寒草甸是青藏高原广泛分布的植被类型之一,面积约120万km2,地处青藏高原腹地的当雄草原站即位于该类植被的典型分布区。以2003年8~10月中旬在该站用涡度相关法连续观测的CO2通量数据资料为基础,分析了高寒草甸生态系统8~10月份净二氧化碳交换量(NEE)的日变化规律,及其与光合有效辐射、降水、温度等环境因子之间的关系。结果表明,8~10月份的日均NEE有明显的日变化,表现为单峰型,通常在地方时11:00~12:00左右达到碳吸收的最大值,平均为-0.2680mgCO2/(m2·s)(-6.0800μmolCO2/(m2·s))。白天的NEE与光合有效辐射之间符合很好的直角双曲线关系,表观量子产额平均为0.0203μmolCO2/μmolPAR,表观最大光合速率平均为9.7411μmolCO2/(m2·s)。夜晚的NEE与5cm地温有很好的指数函数关系。     

增温对青藏高原高寒草甸生态系统固碳通量影响的模拟研究

低温被广泛认为是高寒草甸生态系统首要限制性因子,因此增温可能会在某种程度上促进初级生产力,但是也可能由于土壤水分、N素营养状况的改变形成新胁迫而抑制生产力提高。此外,生态系统呼吸由于增温而提高的幅度也可能高于初级生产力提高的幅度,造成总碳库平衡的改变。利用青藏高原海北高寒草甸实测数据对生态系统过程模型Biome-BGC(V.4.2)进行了参数化,并利用研究区实测土壤水分(0-40 cm)和其它观测数据对模型进行了检验,证明模型模拟结果较为可靠。模型使用2005-2008年的海北气象站实测气象数据包括气温、降水等作为驱动数据,模拟了增温1.2-1.7℃下青藏高原海北定位站高寒草甸生态系统碳通量的变化,并整合分析增温试验平台上已发表的试验,与模拟结果进行对比,探讨增温对海北高寒草甸生态系统碳收支的可能影响。结果表明:2005-2008年青藏高原高寒草甸生态系统为弱的碳汇,短期增温导致系统净碳固定增加。增温直接影响系统碳通量,也通过土壤水分和土壤矿化氮变化间接影响碳通量,相比土壤水分和氮素,增温对影响碳通量变化过程中的效应更大;研究也揭示,在增温条件下,植物对土壤矿化氮的吸收量小于有机质分解产生的土壤矿化氮量,土壤矿化氮含量增加。     

Seasonal patterns of root and shoot interactions in an alpine meadow on the Tibetan Plateau

Aims Numerous studies have showed that the balance between negative and positive plant–plant interactions shifted along environmental gradients. But little is known how the positive or negative plant–plant interactions varied with temporal fluctuating habitat conditions and plant ontogenetic phases.Methods In a 2-year experiment, the four perennial grasses (Kobresia humilis, Stipa aliena, Elymus nutans and Saussurea superba) were grown under four interaction treatments (no root or shoot interaction, only shoot interaction, only root interaction, root and shoot interaction). Intensity of above- and belowground interactions is proposed to vary with the fluctuation of seasonal climatic conditions and soil available nutrients. Here we report measurements of above- and belowground interactions during entire growing season. Correlation between plant interaction intensity and seasonal soil available N as well as habitat climate conditions was also performed.Important findings Our experiment found that root interactions had negative effect on plant growth for the four species during growing season. However, both negative and positive shoot interactions occurred among the four species. Despite there being shoot facilitative effect for E. nutans and S. superba, the full interaction was negative, suggested that root interaction take more important role on plant growth than that of shoot interaction. The interaction between root and shoot effect varied as a function of species identity and growth phases. The weak correlation of plant interaction intensity to habitat environmental factors suggested that plant ontogenetic characteristics may be primary factors causing temporal variation in plant interaction.