不同草地类型土壤丛枝菌根真菌群落特征及其驱动因素

doi:10.17521/cjpe.2023.0244

摘要/Abstract

摘要：

丛枝菌根真菌(AMF)在草地生态系统中广泛分布, 对维持草地生态系统生产力和稳定性具有重要意义, 而中国天然草地生态系统AMF群落的分布特征及其驱动机制尚不明确。该研究以常见的18种草地类型为研究对象, 通过野外调查不同植物群落特征, 分析土壤理化性质和AMF群落特征, 探究中国草地生态系统不同草地类型土壤AMF群落特征及其驱动因素。结果表明, 近明球囊霉属(Claroideoglomus)、隔球囊霉属(Septoglomus)、多孢囊霉属(Diversispora)、根孢囊霉属(Rhizophagus)、无梗囊霉属(Acaulospora)、球囊霉属(Glomus)和双型囊霉属(Ambispora)在不同草地类型AMF群落中占据优势地位(相对多度>15%)。不同草地类型AMF群落在物种组成和多样性上存在差异, 且紫花针茅(Stipa purpurea)草地AMF群落多样性和稀有可操作分类单元(OTU) (相对多度<0.1%的OTU)占比相对较高。结构方程模型表明, AMF群落多样性受植物群落多样性和土壤pH的显著影响, 而AMF群落组成受温度、土壤含水率和植物群落组成的显著影响。综上所述, 中国天然草地AMF资源丰富, 且不同草地类型AMF群落存在较大差异。草地生态系统植物群落特征对AMF群落特征具有强烈的驱动作用, 保护植物多样性对于维持土壤微生物群落稳定性具有重要意义。

关键词: 丛枝菌根真菌, 草地生态系统, 群落组成, 多样性, 驱动因素

Abstract:

Aims Arbuscular mycorrhizal fungi (AMF) are widely distributed in grassland ecosystems. They make great contribution to the productivity and stability of grassland ecosystems. Our purpose was to explore the AMF community distribution characteristics and driving mechanism in grassland ecosystems.

Methods This study was conducted in 18 different grassland types. We demonstrate the AMF community characteristics and driving factors by investigating plant community characteristics, analyzing soil physicochemical properties and AMF communities.

Important findings In different grassland types, the dominant genera of AMF community were Claroideoglomus, Septoglomus, Diversispora, Rhizophagus, Acaulospora, Glomus and Ambispora. In addition, there exist significant differences in speicies composition and diversity of AMF communities, and the AMF diversity and rare operational taxonomic unites (OTUs) in Stipa purpurea grassland was relatively higher than those of other grassland types. Results from the structural equation model showed that AMF community diversity was significantly influenced by plant community diversity and soil pH, and AMF community composition was significantly influenced by temperature, soil moisture content and plant community composition. To sum up, abundant AMF resources were contained within Chinese natural grassland, and the AMF community characteristics in different grassland types are significantly different. Meanwhile, plant communities have remarkable driving effect on AMF communities in grassland ecosystems. So protecting grassland plant diversity is of great importance for maintaining soil microbial community stability.

Key words: arbuscular mycorrhizal fungi, grassland ecosystem, community composition, diversity, driving factor

陈科宇, 邢森, 唐玉, 孙佳慧, 任世杰, 张静, 纪宝明. 不同草地类型土壤丛枝菌根真菌群落特征及其驱动因素. 植物生态学报, 2024, 48(5): 660-674. DOI: 10.17521/cjpe.2023.0244

CHEN Ke-Yu, XING Sen, TANG Yu, SUN Jia-Hui, REN Shi-Jie, ZHANG Jing, JI Bao-Ming. Arbuscular mycorrhizal fungal community characteristics and driving factors in different grassland types. Chinese Journal of Plant Ecology, 2024, 48(5): 660-674. DOI: 10.17521/cjpe.2023.0244

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0244

https://www.plant-ecology.com/CN/Y2024/V48/I5/660

图/表 8

参考文献 66

[1]	Aguilera P, Cornejo P, Borie F, Barea JM, von Baer E, Oehl F (2014). Diversity of arbuscular mycorrhizal fungi associated with Triticum aestivum L. plants growing in an Andosol with high aluminum level. Agriculture, Ecosystems & Environment, 186, 178-184.
[2]	Baar J (2010). Restoration of plant communities in the Netherlands through the application of arbuscular mycorrhizal fungi. Symbiosis, 52, 87-94.
[3]	Bainard LD, Chagnon PL, Cade-Menun BJ, Lamb EG, LaForge K, Schellenberg M, Hamel C (2017). Plant communities and soil properties mediate agricultural land use impacts on arbuscular mycorrhizal fungi in the mixed prairie ecoregion of the North American Great Plains. Agriculture, Ecosystems & Environment, 249, 187-195.
[4]	Barceló M, van Bodegom PM, Soudzilovskaia NA (2019). Climate drives the spatial distribution of mycorrhizal host plants in terrestrial ecosystems. Journal of Ecology, 107, 2564-2573.
[5]	Bever JD (2003). Soil community feedback and the coexistence of competitors: conceptual frameworks and empirical tests. New Phytologist, 157, 465-473. DOI PMID
[6]	Bever JD, Platt TG, Morton ER (2012). Microbial population and community dynamics on plant roots and their feedbacks on plant communities. Annual Review of Microbiology, 66, 265-283. DOI PMID
[7]	Bever JD, Westover KM, Antonovics J (1997). Incorporating the soil community into plant population dynamics: the utility of the feedback approach. Journal of Ecology, 85, 561-573.
[8]	Cavagnaro TR (2016). Soil moisture legacy effects: impacts on soil nutrients, plants and mycorrhizal responsiveness. Soil Biology & Biochemistry, 95, 173-179.
[9]	Chaudhary VB, O’Dell TE, Rillig MC, Johnson NC (2014). Multiscale patterns of arbuscular mycorrhizal fungal abundance and diversity in semiarid shrublands. Fungal Ecology, 12, 32-43.
[10]	Chen KY, Zhang J, Muneer MA, Xue K, Niu HS, Ji BM (2023). Plant community and soil available nutrients drive arbuscular mycorrhizal fungal community shifts during alpine meadow degradation. Fungal Ecology, 62, 101211. DOI: 10.1016/j.funeco.2022.101211.
[11]	Davison J, García de León D, Zobel M, Moora M, Bueno CG, Barceló M, Gerz M, León D, Meng YM, Pillar VD, Sepp SK, Soudzilovaskaia NA, Tedersoo L, Vaessen S, et al. (2020). Plant functional groups associate with distinct arbuscular mycorrhizal fungal communities. New Phytologist, 226, 1117-1128. DOI PMID
[12]	Davison J, Moora M, Öpik M, Adholeya A, Ainsaar L, Bâ A, Burla S, Diedhiou AG, Hiiesalu I, Jairus T, Johnson NC, Kane A, Koorem K, Kochar M, Ndiaye C, et al. (2015). Global assessment of arbuscular mycorrhizal fungus diversity reveals very low endemism. Science, 349, 970-973. DOI PMID
[13]	de la Sota-Ricaldi AM, Rengifo del Águila S, Blas Sevillano R, López-García Á, Corazon-Guivin MA (2023). Beta diversity of arbuscular mycorrhizal communities increases in time after crop establishment of Peruvian Sacha inchi (Plukenetia volubilis). Journal of Fungi, 9, 194. DOI: 10.3390/jof9020194.
[14]	Elliott AJ, Daniell TJ, Cameron DD, Field KJ (2021). A commercial arbuscular mycorrhizal inoculum increases root colonization across wheat cultivars but does not increase assimilation of mycorrhiza-acquired nutrients. Plants, People, Planet, 3, 588-599. DOI PMID
[15]	Fei SL, Kivlin SN, Domke GM, Jo I, LaRue EA, Phillips RP (2022). Coupling of plant and mycorrhizal fungal diversity: its occurrence, relevance, and possible implications under global change. New Phytologist, 234, 1960-1966. DOI PMID
[16]	Frew A, Heuck MK, Aguilar-Trigueros CA (2023). Host filtering, not competitive exclusion, may be the main driver of arbuscular mycorrhizal fungal community assembly under high phosphorus. Functional Ecology, 37, 1856-1869.
[17]	Fuchs B, Haselwandter K (2004). Red list plants: colonization by arbuscular mycorrhizal fungi and dark septate endophytes. Mycorrhiza, 14, 277-281. PMID
[18]	Gao C, Kim YC, Zheng Y, Yang W, Chen L, Ji NN, Wan SQ, Guo LD (2016). Increased precipitation, rather than warming, exerts a strong influence on arbuscular mycorrhizal fungal community in a semiarid steppe ecosystem. Botany, 94, 459-469.
[19]	Guo X, Wang Z, Zhang J, Wang P, Li YM, Ji BM (2021). Host-specific effects of arbuscular mycorrhizal fungi on two Caragana species in desert grassland. Journal of Fungi, 7, 1077. DOI: 10.3390/jof7121077.
[20]	He YJ, Jiang CH, Yang H, Wang YJ, Zhong ZC (2017). Arbuscular mycorrhizal fungal composition affects the growth and nutrient acquisition of two plants from a karst area. Sains Malaysiana, 46, 1701-1708.
[21]	Jamiołkowska A, Księżniak A, Gałązka A, Hetman B, Kopacki M, Skwaryło-Bednarz B (2018). Impact of abiotic factors on development of the community of arbuscular mycorrhizal fungi in the soil: a review. International Agrophysics, 32, 133-140.
[22]	Ji BM, Bentivenga SP, Casper BB (2010). Evidence for ecological matching of whole AM fungal communities to the local plant-soil environment. Ecology, 91, 3037-3046. PMID
[23]	Ji BM, Bever JD (2016). Plant preferential allocation and fungal reward decline with soil phosphorus: implications for mycorrhizal mutualism. Ecosphere, 7, e01256. DOI: 10.1002/ecs2.1256.
[24]	Ji MK, Kong WD, Stegen J, Yue LY, Wang F, Dong XB, Cowan DA, Ferrari BC (2020). Distinct assembly mechanisms underlie similar biogeographical patterns of rare and abundant bacteria in Tibetan Plateau grassland soils. Environmental Microbiology, 22, 2261-2272. DOI PMID
[25]	Johnson D, Martin F, Cairney JWG, Anderson IC (2012). The importance of individuals: intraspecific diversity of mycorrhizal plants and fungi in ecosystems. New Phytologist, 194, 614-628. DOI PMID
[26]	Kokkoris V, Li YL, Hamel C, Hanson K, Hart M (2019). Site specificity in establishment of a commercial arbuscular mycorrhizal fungal inoculant. Science of the Total Environment, 660, 1135-1143. DOI
[27]	Koziol L, Bever JD (2019). Mycorrhizal feedbacks generate positive frequency dependence accelerating grassland succession. Journal of Ecology, 107, 622-632. DOI
[28]	Kytöviita MM, Ruotsalainen AL (2007). Mycorrhizal benefit in two low arctic herbs increases with increasing temperature. American Journal of Botany, 94, 1309-1315.
[29]	Lee J, Lee SS, Young JPW (2008). Improved PCR primers for the detection and identification of arbuscular mycorrhizal fungi. FEMS Microbiology Ecology, 65, 339-349. DOI PMID
[30]	Lenoir I, Fontaine J, Lounès-Hadj Sahraoui A (2016). Arbuscular mycorrhizal fungal responses to abiotic stresses: a review. Phytochemistry, 123, 4-15. DOI PMID
[31]	Liu M, Zheng R, Bai SL, Bai Y, Wang JG (2017). Slope aspect influences arbuscular mycorrhizal fungus communities in arid ecosystems of the Daqingshan Mountains, Inner Mongolia, North China. Mycorrhiza, 27, 189-200. DOI PMID
[32]	Liu YJ, He JX, Shi GX, An LZ, Öpik M, Feng HY (2011). Diverse communities of arbuscular mycorrhizal fungi inhabit sites with very high altitude in Tibet Plateau. FEMS Microbiology Ecology, 78, 355-365. DOI PMID
[33]	Liu YJ, Shi GX, Mao L, Cheng G, Jiang SJ, Ma XJ, An LZ, Du GZ, Johnson NC, Feng HY (2012). Direct and indirect influences of 8 yr of nitrogen and phosphorus fertilization on Glomeromycota in an alpine meadow ecosystem. New Phytologist, 194, 523-535. DOI PMID
[34]	López-García Á, Palenzuela J, Barea JM, Azcón-Aguilar C (2014). Life-history strategies of arbuscular mycorrhizal fungi determine succession into roots of Rosmarinus officinalis L., a characteristic woody perennial plant species from Mediterranean ecosystems. Plant and Soil, 379, 247-260.
[35]	Lu XL, Lu XN, Liao YC (2018). Effect of tillage treatment on the diversity of soil arbuscular mycorrhizal fungal and soil aggregate-associated carbon content. Frontiers in Microbiology, 9, 2986. DOI: 10.3389/fmicb.2018.02986.
[36]	Luan L, Jiang YJ, Dini-Andreote F, Crowther TW, Li PF, Bahram M, Zheng J, Xu QS, Zhang XX, Sun B (2023). Integrating pH into the metabolic theory of ecology to predict bacterial diversity in soil. Proceedings of the National Academy of Sciences of the United States of America, 120, e2207832120. DOI: 10.1073/pnas.2207832120.
[37]	Luo X, Su XL, Cui JM, Lou Y, Li R, Luo XM, Zeng YQ, Xu YB, Dong JY (2016). Biodiversity of arbuscular mycorrhizal fungi in the drawdown zone of the Three Gorges Reservoir under different fertilization histories. Ecological Research, 31, 407-416.
[38]	Luo YF, Wang ZK, He YL, Li GF, Lv XH, Zhuang L (2020). High-throughput sequencing analysis of the rhizosphere arbuscular mycorrhizal fungi (AMF) community composition associated with Ferula sinkiangensis. BMC Microbiology, 20, 335. DOI: 10.1186/s12866-020-02024-x.
[39]	Ma XC, Xu X, Geng QH, Luo YQ, Ju CH, Li Q, Zhou Y (2023). Global arbuscular mycorrhizal fungal diversity and abundance decreases with soil available phosphorus. Global Ecology and Biogeography, 32, 1423-1434.
[40]	Martínez-García LB, Richardson SJ, Tylianakis JM, Peltzer DA, Dickie IA (2015). Host identity is a dominant driver of mycorrhizal fungal community composition during ecosystem development. New Phytologist, 205, 1565-1576. DOI PMID
[41]	Menzel A, Hempel S, Manceur AM, Götzenberger L, Moora M, Rillig MC, Zobel M, Kühn I (2016). Distribution patterns of arbuscular mycorrhizal and non-mycorrhizal plant species in Germany. Perspectives in Plant Ecology, Evolution and Systematics, 21, 78-88.
[42]	Muneer MA, Wang P, Zhang J, Li YM, Munir MZ, Ji BM (2020). Formation of common mycorrhizal networks significantly affects plant biomass and soil properties of the neighboring plants under various nitrogen levels. Microrganisms, 8, 230. DOI: 10.3390/microorganisms8020230.
[43]	Öpik M, Zobel M, Cantero JJ, Davison J, Facelli JM, Hiiesalu I, Jairus T, Kalwij JM, Koorem K, Leal ME, Liira J, Metsis M, Neshataeva V, Paal J, Phosri C, et al. (2013). Global sampling of plant roots expands the described molecular diversity of arbuscular mycorrhizal fungi. Mycorrhiza, 23, 411-430. DOI PMID
[44]	Powell JR, Rillig MC (2018). Biodiversity of arbuscular mycorrhizal fungi and ecosystem function. New Phytologist, 220, 1059-1075. DOI PMID
[45]	Ramana JV, Tylianakis JM, Ridgway HJ, Dickie IA (2023). Root diameter, host specificity and arbuscular mycorrhizal fungal community composition among native and exotic plant species. New Phytologist, 239, 301-310.
[46]	Sarr PS, Sugiyama A, Begoude ADB, Yazaki K, Araki S, Nawata E (2019). Diversity and distribution of Arbuscular Mycorrhizal Fungi in cassava (Manihot esculenta Crantz) croplands in Cameroon as revealed by Illumina MiSeq. Rhizosphere, 10, 100147. DOI: 10.1016/j.rhisph.2019.100147.
[47]	Sato K, Suyama Y, Saito M, Sugawara K (2005). A new primer for discrimination of arbuscular mycorrhizal fungi with polymerase chain reaction-denature gradient gel electrophoresis. Grassland Science, 51, 179-181.
[48]	Staddon PL, Thompson K, Jakobsen I, Grime JP, Askew AP, Fitter AH (2003). Mycorrhizal fungal abundance is affected by long-term climatic manipulations in the field. Global Change Biology, 9, 186-194.
[49]	Sun XF, Su YY, Zhang Y, Wu MY, Zhang Z, Pei KQ, Sun LF, Wan SQ, Liang Y (2013). Diversity of arbuscular mycorrhizal fungal spore communities and its relations to plants under increased temperature and precipitation in a natural grassland. Chinese Science Bulletin, 58, 4109-4119.
[50]	Thonar C, Frossard E, Šmilauer P, Jansa J (2014). Competition and facilitation in synthetic communities of arbuscular mycorrhizal fungi. Molecular Ecology, 23, 733-746. DOI PMID
[51]	Urcelay C, Díaz S (2003). The mycorrhizal dependence of subordinates determines the effect of arbuscular mycorrhizal fungi on plant diversity. Ecology Letters, 6, 388-391.
[52]	van Geel M, De Beenhouwer M, Lievens B, Honnay O (2016). Crop-specific and single-species mycorrhizal inoculation is the best approach to improve crop growth in controlled environments. Agronomy for Sustainable Development, 36, 37. DOI: 10.1007/s13593-016-0373-y.
[53]	Wagg C, Bender SF, Widmer F, van der Heijden MGA (2014). Soil biodiversity and soil community composition determine ecosystem multifunctionality. Proceedings of the National Academy of Sciences of the United States of America, 111, 5266-5270. DOI PMID
[54]	Wagg C, Jansa J, Stadler M, Schmid B, van der Heijden MGA (2011). Mycorrhizal fungal identity and diversity relaxes plant-plant competition. Ecology, 92, 1303-1313. PMID
[55]	Wang B, Qiu YL (2006). Phylogenetic distribution and evolution of mycorrhizas in land plants. Mycorrhiza, 16, 299-363. PMID
[56]	Wang GM, Stribley DP, Tinker PB, Walker C (1993). Effects of pH on arbuscular mycorrhiza I. Field observations on the long-term liming experiments at Rothamsted and Woburn. New Phytologist, 124, 465-472.
[57]	Wang J, Yang CX, Sun XM, Zhang HO, Guo Z, Cao TT, Li J (2021). Growth responses of Sorghum bicolor (L.) Moench to arbuscular mycorrhizal fungi under simulated nitrogen deposition. Bangladesh Journal of Botany, 50, 933-938.
[58]	Wu E, Li ZX, Zhang Y, Wang CM, Yun JF (2009). Effect of grassland degradation on arbuscular mycorrhizal symbiosis of Leymus chinensis (Trin.) Tzvel. in typical steppe. Acta Agrestia Sinica, 17, 731-734.
	[乌恩, 李重祥, 张宇, 王聪明, 云锦凤 (2009). 草原退化对典型草原羊草菌根共生的影响. 草地学报, 17, 731-734.] DOI
[59]	Wu H, Yang J, Fu W, Rillig MC, Cao Z, Zhao A, Hao Z, Zhang X, Chen B, Han X (2023). Identifying thresholds of nitrogen enrichment for substantial shifts in arbuscular mycorrhizal fungal community metrics in a temperate grassland of Northern China. New Phytologist, 237, 279-294.
[60]	Xiao D, Chen Y, He XY, Xu ZH, Hosseini Bai S, Zhang W, Cheng M, Hu PL, Wang KL (2021). Temperature and precipitation significantly influence the interactions between arbuscular mycorrhizal fungi and diazotrophs in karst ecosystems. Forest Ecology and Management, 497, 119464. DOI: 10.1016/j.foreco.2021.119464.
[61]	Xiao D, Liu X, Yang R, Tan YJ, Zhang W, Deng CX, He XY, Wang KL (2020). Nitrogen input through chemical and biological pathway affects arbuscular mycorrhizal fungal communities in forage grass cultivation in southwest China. Applied Soil Ecology, 150, 103454. DOI: 10.1016/j.apsoil.2019.103454.
[62]	Yang L, Liu Y, Sui X, Zhang T, Xu N, Ni H, Wu Y (2018). Effects of different land uses on community structure of soil arbuscular mycorrhizal fungi in sanjiang wetlands. Bangladesh Journal of Botany, 47, 689-697.
[63]	Ye XH, Li JQ, Ma JH, Fan QF, Yu N, Zhang YL, Zou HT, Zhang YL (2023). Drip irrigation at a soil water suction of 30 kPa helps AMF and GRSP to enhance greenhouse macro-aggregates. Agronomy, 13, 522. DOI: 10.3390/agronomy13020522.
[64]	Zhang L, Zhou J, George TS, Limpens E, Feng G (2022). Arbuscular mycorrhizal fungi conducting the hyphosphere bacterial orchestra. Trends in Plant Science, 27, 402-411.
[65]	Zhong R, Xia C, Ju YW, Zhang XX, Duan TY, Nan ZB, Li CJ (2021). A foliar Epichloë endophyte and soil moisture modified belowground arbuscular mycorrhizal fungal biodiversity associated with Achnatherum inebrians. Plant and Soil, 458, 105-122.
[66]	Zhou ZH, Wang CK, Luo YQ (2020). Meta-analysis of the impacts of global change factors on soil microbial diversity and functionality. Nature Communications, 11, 3072. DOI: 10.1038/s41467-020-16881-7.

草地类型 Grassland type	经度 Longitude (° E)	纬度 Latitude (° N)	海拔 Altitude (m)	1-8月平均气温 Average air temperature of Jan. to Aug. (℃)	1-8月降水量 Precipitation of Jan. to Aug. (mm)
高山嵩草-垂穗披碱草草地 Carex parvula - Elymus nutans grassland	91.90	29.72	4 056	-4.29	576.73
高山嵩草-圆穗蓼草地 C. parvula - Bistorta macrophylla grassland	93.25	30.69	4 473	-9.73	328.00
蕨麻草地 Argentina anserina grassland	91.15	31.12	4 488	-2.51	316.85
高山早熟禾草地 Poa alpina grassland	91.90	29.73	4 050	-4.29	576.73
西藏嵩草草地 C. tibetikobresia grassland	93.23	31.92	4 543	-9.99	478.53
垂穗披碱草草地 E. nutans grassland	98.27	30.87	3 640	-4.42	579.03
火绒草草地 Leontopodium leontopodioides grassland	91.51	31.87	4 502	-3.21	392.97
紫花针茅草地 Stipa purpurea grassland	90.32	31.39	4 641	-2.54	340.49
紫花针茅-青藏薹草草地 S. purpurea - C. moorcroftii grassland	84.83	32.00	4 578	-1.43	92.64
沙生薹草草地 C. praeclara grassland	84.03	29.78	4 816	-8.25	186.31
丝颖针茅草地 S. capillacea grassland	84.03	29.77	4 836	-8.22	186.31
大花嵩草草地 C. nudicarpa grassland	88.47	28.21	4 700	0.44	271.86
羊草-短花针茅草地 Leymus chinensis - S. breviflora grassland	111.22	41.25	1 701	4.40	96.12
短花针茅-冷蒿草地 S. breviflora - Artemisia frigida grassland	111.22	41.26	1 690	4.39	95.89
羊草-糙隐子草草地 L. chinensis - Cleistogenes squarrosa grassland	116.34	44.27	1 075	3.00	136.15
大针茅草地 S. grandis grassland	116.44	43.48	1 377	2.71	132.78
羊草-狼针草草地 L. chinensis - S. baicalensis grassland	120.00	49.34	613	-0.18	116.59
狼针草草地 S. baicalensis grassland	119.98	49.60	644	-0.31	119.26

草地类型 Grassland type	经度 Longitude (° E)	纬度 Latitude (° N)	海拔 Altitude (m)	1-8月平均气温 Average air temperature of Jan. to Aug. (℃)	1-8月降水量 Precipitation of Jan. to Aug. (mm)
高山嵩草-垂穗披碱草草地 Carex parvula - Elymus nutans grassland	91.90	29.72	4 056	-4.29	576.73
高山嵩草-圆穗蓼草地 C. parvula - Bistorta macrophylla grassland	93.25	30.69	4 473	-9.73	328.00
蕨麻草地 Argentina anserina grassland	91.15	31.12	4 488	-2.51	316.85
高山早熟禾草地 Poa alpina grassland	91.90	29.73	4 050	-4.29	576.73
西藏嵩草草地 C. tibetikobresia grassland	93.23	31.92	4 543	-9.99	478.53
垂穗披碱草草地 E. nutans grassland	98.27	30.87	3 640	-4.42	579.03
火绒草草地 Leontopodium leontopodioides grassland	91.51	31.87	4 502	-3.21	392.97
紫花针茅草地 Stipa purpurea grassland	90.32	31.39	4 641	-2.54	340.49
紫花针茅-青藏薹草草地 S. purpurea - C. moorcroftii grassland	84.83	32.00	4 578	-1.43	92.64
沙生薹草草地 C. praeclara grassland	84.03	29.78	4 816	-8.25	186.31
丝颖针茅草地 S. capillacea grassland	84.03	29.77	4 836	-8.22	186.31
大花嵩草草地 C. nudicarpa grassland	88.47	28.21	4 700	0.44	271.86
羊草-短花针茅草地 Leymus chinensis - S. breviflora grassland	111.22	41.25	1 701	4.40	96.12
短花针茅-冷蒿草地 S. breviflora - Artemisia frigida grassland	111.22	41.26	1 690	4.39	95.89
羊草-糙隐子草草地 L. chinensis - Cleistogenes squarrosa grassland	116.34	44.27	1 075	3.00	136.15
大针茅草地 S. grandis grassland	116.44	43.48	1 377	2.71	132.78
羊草-狼针草草地 L. chinensis - S. baicalensis grassland	120.00	49.34	613	-0.18	116.59
狼针草草地 S. baicalensis grassland	119.98	49.60	644	-0.31	119.26

草地类型 Grassland type	盖度 Coverage	高度 Height	丰富度 Richness	Simpson多样性指数 Simpson diversity index	Shannon-Wiener多样性指数 Shannon-Wiener diversity index	Pielou均匀度指数 Pielou evenness index
高山嵩草-垂穗披碱草草地 Carex parvula - Elymus nutans grassland	98.40 ± 0.40^a	26.51 ± 1.46^c	9.20 ± 0.73^c	0.86 ± 0.01^ab	2.07 ± 0.05^cd	0.94 ± 0.01^ab
高山嵩草-圆穗蓼草地 C. parvula - Bistorta macrophylla grassland	98.00 ± 0.45^a	7.53 ± 0.26^hij	13.40 ± 0.51^a	0.89 ± 0.01^a	2.42 ± 0.02^a	0.94 ± 0.01^ab
蕨麻草地 Argentina anserina grassland	48.00 ± 1.64^e	7.01 ± 0.71^ij	2.80 ± 0.37^f	0.51 ± 0.04^hi	0.82 ± 0.10^jk	0.89 ± 0.02^abcd
高山早熟禾草地 Poa alpina grassland	94.20 ± 1.66^ab	12.01 ± 0.78^efg	12.40 ± 0.40^ab	0.89 ± 0.01^a	2.33 ± 0.03^ab	0.94 ± 0.02^ab
西藏嵩草草地 C. tibetikobresia grassland	95.60 ± 1.50^ab	4.71 ± 0.23^j	11.60 ± 1.03^b	0.85 ± 0.02^ab	2.19 ± 0.08^bc	0.90 ± 0.01^abcd
垂穗披碱草草地 E. nutans grassland	96.60 ± 0.51^ab	23.57 ± 1.62^cd	8.60 ± 0.51^c	0.84 ± 0.02^abc	1.87 ± 0.05^def	0.87 ± 0.01^bcde
火绒草草地 Leontopodium leontopodioides grassland	64.20 ± 2.29^d	10.55 ± 0.57^fgh	3.60 ± 0.24^f	0.59 ± 0.03^fg	1.08 ± 0.06^hi	0.90 ± 0.03^abcd
紫花针茅草地 Stipa purpurea grassland	48.00 ± 0.95^e	12.63 ± 0.18^efg	7.00 ± 0.45^d	0.78 ± 0.01^bcd	1.75 ± 0.05^efg	0.90 ± 0.01^abcd
紫花针茅-青藏薹草草地 S. purpurea - C. moorcroftii grassland	43.00 ± 3.74^ef	14.64 ± 0.21^e	3.00 ± 0.32^f	0.58 ± 0.01^gh	0.98 ± 0.01^hij	0.89 ± 0.01^abcd
沙生薹草草地 C. praeclara grassland	40.40 ± 1.44^fg	9.84 ± 0.74^ghi	3.40 ± 0.24^f	0.73 ± 0.06^de	1.00 ± 0.02 ^hij	0.85 ± 0.06^de
丝颖针茅草地 S. capillacea grassland	36.00 ± 1.30^g	13.91 ± 0.54^ef	3.00 ± 0.45^f	0.67 ± 0.02^ef	0.88 ± 0.16^ijk	0.81 ± 0.04^e
大花嵩草草地 C. nudicarpa grassland	29.00 ± 1.38^h	9.51 ± 1.28^ghi	2.40 ± 0.24^f	0.49 ± 0.03ⁱ	0.75 ± 0.07^k	0.91 ± 0.02^abcd
羊草-短花针茅草地 Leymus chinensis - S. breviflora grassland	43.40 ± 3.33^ef	20.44 ± 0.52^d	3.40 ± 0.24^f	0.67 ± 0.01^ef	1.15 ± 0.05^h	0.95 ± 0.01^a
短花针茅-冷蒿草地 S. breviflora - Artemisia frigida grassland	45.00 ± 1.70^ef	25.54 ± 1.35^c	3.00 ± 0.45^f	0.52 ± 0.05^ghi	0.90 ± 0.13^ijk	0.85 ± 0.01^cde
羊草-糙隐子草草地 L. chinensis - Cleistogenes squarrosa grassland	79.00 ± 2.63^c	31.59 ± 0.68^b	6.40 ± 0.24^de	0.77 ± 0.01^cd	1.65 ± 0.04^g	0.88 ± 0.01^abcde
大针茅草地 S. grandis grassland	65.60 ± 0.87^d	21.84 ± 1.40^d	5.40 ± 0.24^e	0.76 ± 0.01^cd	1.55 ± 0.03^g	0.92 ± 0.01^abcd
羊草-狼针草草地 L. chinensis - S. baicalensis grassland	95.20 ± 2.65^ab	36.32 ± 2.86^a	8.40 ± 0.24^c	0.83 ± 0.01^abc	1.93 ± 0.03^de	0.91 ± 0.01^abcd
狼针草草地 S. baicalensis grassland	91.20 ± 1.20^b	34.12 ± 0.46^ab	6.40 ± 0.68^de	0.79 ± 0.01^bcd	1.69 ± 0.09^fg	0.92 ± 0.01^abc

草地类型 Grassland type	盖度 Coverage	高度 Height	丰富度 Richness	Simpson多样性指数 Simpson diversity index	Shannon-Wiener多样性指数 Shannon-Wiener diversity index	Pielou均匀度指数 Pielou evenness index
高山嵩草-垂穗披碱草草地 Carex parvula - Elymus nutans grassland	98.40 ± 0.40^a	26.51 ± 1.46^c	9.20 ± 0.73^c	0.86 ± 0.01^ab	2.07 ± 0.05^cd	0.94 ± 0.01^ab
高山嵩草-圆穗蓼草地 C. parvula - Bistorta macrophylla grassland	98.00 ± 0.45^a	7.53 ± 0.26^hij	13.40 ± 0.51^a	0.89 ± 0.01^a	2.42 ± 0.02^a	0.94 ± 0.01^ab
蕨麻草地 Argentina anserina grassland	48.00 ± 1.64^e	7.01 ± 0.71^ij	2.80 ± 0.37^f	0.51 ± 0.04^hi	0.82 ± 0.10^jk	0.89 ± 0.02^abcd
高山早熟禾草地 Poa alpina grassland	94.20 ± 1.66^ab	12.01 ± 0.78^efg	12.40 ± 0.40^ab	0.89 ± 0.01^a	2.33 ± 0.03^ab	0.94 ± 0.02^ab
西藏嵩草草地 C. tibetikobresia grassland	95.60 ± 1.50^ab	4.71 ± 0.23^j	11.60 ± 1.03^b	0.85 ± 0.02^ab	2.19 ± 0.08^bc	0.90 ± 0.01^abcd
垂穗披碱草草地 E. nutans grassland	96.60 ± 0.51^ab	23.57 ± 1.62^cd	8.60 ± 0.51^c	0.84 ± 0.02^abc	1.87 ± 0.05^def	0.87 ± 0.01^bcde
火绒草草地 Leontopodium leontopodioides grassland	64.20 ± 2.29^d	10.55 ± 0.57^fgh	3.60 ± 0.24^f	0.59 ± 0.03^fg	1.08 ± 0.06^hi	0.90 ± 0.03^abcd
紫花针茅草地 Stipa purpurea grassland	48.00 ± 0.95^e	12.63 ± 0.18^efg	7.00 ± 0.45^d	0.78 ± 0.01^bcd	1.75 ± 0.05^efg	0.90 ± 0.01^abcd
紫花针茅-青藏薹草草地 S. purpurea - C. moorcroftii grassland	43.00 ± 3.74^ef	14.64 ± 0.21^e	3.00 ± 0.32^f	0.58 ± 0.01^gh	0.98 ± 0.01^hij	0.89 ± 0.01^abcd
沙生薹草草地 C. praeclara grassland	40.40 ± 1.44^fg	9.84 ± 0.74^ghi	3.40 ± 0.24^f	0.73 ± 0.06^de	1.00 ± 0.02 ^hij	0.85 ± 0.06^de
丝颖针茅草地 S. capillacea grassland	36.00 ± 1.30^g	13.91 ± 0.54^ef	3.00 ± 0.45^f	0.67 ± 0.02^ef	0.88 ± 0.16^ijk	0.81 ± 0.04^e
大花嵩草草地 C. nudicarpa grassland	29.00 ± 1.38^h	9.51 ± 1.28^ghi	2.40 ± 0.24^f	0.49 ± 0.03ⁱ	0.75 ± 0.07^k	0.91 ± 0.02^abcd
羊草-短花针茅草地 Leymus chinensis - S. breviflora grassland	43.40 ± 3.33^ef	20.44 ± 0.52^d	3.40 ± 0.24^f	0.67 ± 0.01^ef	1.15 ± 0.05^h	0.95 ± 0.01^a
短花针茅-冷蒿草地 S. breviflora - Artemisia frigida grassland	45.00 ± 1.70^ef	25.54 ± 1.35^c	3.00 ± 0.45^f	0.52 ± 0.05^ghi	0.90 ± 0.13^ijk	0.85 ± 0.01^cde
羊草-糙隐子草草地 L. chinensis - Cleistogenes squarrosa grassland	79.00 ± 2.63^c	31.59 ± 0.68^b	6.40 ± 0.24^de	0.77 ± 0.01^cd	1.65 ± 0.04^g	0.88 ± 0.01^abcde
大针茅草地 S. grandis grassland	65.60 ± 0.87^d	21.84 ± 1.40^d	5.40 ± 0.24^e	0.76 ± 0.01^cd	1.55 ± 0.03^g	0.92 ± 0.01^abcd
羊草-狼针草草地 L. chinensis - S. baicalensis grassland	95.20 ± 2.65^ab	36.32 ± 2.86^a	8.40 ± 0.24^c	0.83 ± 0.01^abc	1.93 ± 0.03^de	0.91 ± 0.01^abcd
狼针草草地 S. baicalensis grassland	91.20 ± 1.20^b	34.12 ± 0.46^ab	6.40 ± 0.68^de	0.79 ± 0.01^bcd	1.69 ± 0.09^fg	0.92 ± 0.01^abc

草地类型 Grassland type	TC (g·kg^-1)	TN (g·kg^-1)	TP (g·kg^-1)	NH₄⁺-N (mg·kg^-1)	NO₃^--N(mg·kg^-1)	AP (mg·kg^-1)	pH	土壤含水率 Soil water content (%)
高山嵩草-垂穗披碱草草地 Carex parvula - Elymus nutans grassland	13.70 ± 1.17^def	1.40 ± 0.10^fghi	0.54 ± 0.03^fg	1.80 ± 0.30^c	1.13 ± 0.36^e	0.95 ± 0.14^e	6.47 ± 0.08^e	13.44 ± 1.67^fgh
高山嵩草-圆穗蓼草地 C. parvula - Bistorta macrophylla grassland	52.92 ± 1.15^b	4.57 ± 0.08^c	0.77 ± 0.02^c	22.51 ± 4.72^b	2.53 ± 0.29^de	4.46 ± 0.07^cd	6.23 ± 0.04^ef	34.86 ± 1.22^ab
蕨麻草地 Argentina anserina grassland	59.90 ± 0.47^b	2.94 ± 0.04^de	0.67 ± 0.04^de	2.05 ± 0.29^c	42.20 ± 2.75^bc	4.73 ± 0.18^cd	7.93 ± 0.02^b	20.46 ± 0.47^def
高山早熟禾草地 Poa alpina grassland	59.88 ± 8.07^b	4.94 ± 0.57^bc	0.81 ± 0.04^c	11.15 ± 4.20^c	67.73 ± 36.92^b	8.71 ± 0.58^b	5.97 ± 0.36^fg	31.59 ± 6.07^abc
西藏嵩草草地 C. tibetikobresia grassland	61.66 ± 15.45^b	5.49 ± 1.40^abc	0.73 ± 0.08^cd	10.54 ± 2.40^c	23.58 ± 3.74^cde	8.49 ± 3.88^b	6.25 ± 0.11^ef	33.70 ± 7.89^abc
垂穗披碱草草地 E. nutans grassland	76.54 ± 3.34^a	6.56 ± 0.27^a	1.10 ± 0.01^a	9.34 ± 1.03^c	197.20 ± 5.05^a	9.79 ± 0.11^b	5.79 ± 0.02^g	38.18 ± 1.29^a
火绒草草地 Leontopodium leontopodioides grassland	22.88 ± 0.35^cde	2.04 ± 0.03^defgh	0.56 ± 0.02^f	3.25 ± 0.47^c	6.60 ± 0.76^de	1.98 ± 0.08^de	5.85 ± 0.08^g	16.65 ± 0.70^def
紫花针茅草地 Stipa purpurea grassland	13.98 ± 0.81^def	0.85 ± 0.06^hi	0.37 ± 0.01ⁱ	1.89 ± 0.29^c	7.71 ± 0.49^de	3.44 ± 0.16^de	7.86 ± 0.10^b	4.78 ± 0.91^h
紫花针茅-青藏薹草草地 S. purpurea - C. moorcroftii grassland	18.40 ± 0.41^de	1.89 ± 0.07^efgh	0.22 ± 0.01^j	1.69 ± 0.08^c	5.70 ± 0.30^de	1.74 ± 0.11^de	8.58 ± 0.02^a	4.91 ± 0.67^h
沙生薹草草地 C. praeclara grassland	4.16 ± 0.39^f	0.47 ± 0.02ⁱ	0.26 ± 0.02^j	1.33 ± 0.12^c	1.08 ± 0.17^e	2.83 ± 0.06^de	8.16 ± 0.06^b	11.24 ± 7.10^gh
丝颖针茅草地 S. capillacea grassland	14.70 ± 0.62^def	1.50 ± 0.05^fghi	0.52 ± 0.01^fg	4.05 ± 0.61^c	5.51 ± 0.45^de	4.07 ± 0.09^de	7.05 ± 0.03^cd	4.96 ± 0.84^h
大花嵩草草地 C. nudicarpa grassland	9.06 ± 0.28^ef	1.24 ± 0.03^ghi	0.61 ± 0.01^ef	2.13 ± 0.34^c	3.78 ± 0.42^de	7.22 ± 0.15^bc	8.16 ± 0.06^b	4.30 ± 0.28^h
羊草-短花针茅草地 Leymus chinensis - S. breviflora grassland	18.20 ± 0.30^de	2.17 ± 0.03^defg	0.42 ± 0.01^hi	1.46 ± 0.19^c	32.35 ± 0.99^cd	2.66 ± 0.10^de	7.18 ± 0.02^cd	10.32 ± 0.36^gh
短花针茅-冷蒿草地 S. breviflora - Artemisia frigida grassland	14.18 ± 0.33^def	1.68 ± 0.03^fgh	0.39 ± 0.01ⁱ	2.24 ± 0.44^c	11.51 ± 0.42^de	2.49 ± 0.12^de	6.80 ± 0.05^d	4.22 ± 0.11^h
羊草-糙隐子草草地 L. chinensis - Cleistogenes squarrosa grassland	23.44 ± 1.16^cd	2.54 ± 0.11^def	0.47 ± 0.01^gh	2.80 ± 0.36^c	3.19 ± 0.26^de	8.58 ± 0.45^b	7.27 ± 0.03^c	4.93 ± 0.18^h
大针茅草地 S. grandis grassland	18.32 ± 0.34^de	2.10 ± 0.04^defg	0.37 ± 0.02ⁱ	1.84 ± 0.15^c	4.56 ± 0.29^de	3.36 ± 0.22^de	6.88 ± 0.03^d	5.27 ± 0.18^h
羊草-狼针草草地 L. chinensis - S. baicalensis grassland	34.50 ± 1.40^c	3.14 ± 0.12^d	0.54 ± 0.01^fg	3.58 ± 0.23^c	42.29 ± 3.10^bc	3.24 ± 0.27^de	6.48 ± 0.07^e	26.76 ± 0.93^bcd
狼针草草地 S. baicalensis grassland	61.72 ± 1.81^b	5.77 ± 0.16^ab	0.89 ± 0.02^b	47.99 ± 11.17^a	40.09 ± 5.55^c	16.28 ± 1.32^a	6.54 ± 0.02^e	24.63 ± 0.81^cde