Community characteristics and drivers of arbuscular mycorrhizal fungi in alpine wetlands of the Yellow River Source Region

doi:10.17521/cjpe.2025.0342

Abstract

Abstract:

Aims Investigating the variation in arbuscular mycorrhizal fungal (AMF) community structure and its driving factors in the alpine wetlands of the Yellow River Source Region within Sanjiangyuan National Park can provide a scientific basis for ecosystem conservation and sustainable management in this area.

Methods Alpine riverine wetlands and alpine swamp wetlands in the Yellow River Source Region were selected as the study objects. An integrated approach combining field investigation and sampling, laboratory analyses, and high-throughput sequencing was employed to systematically examine AMF community structure across wetland types and dominant plant species, and to identify the key factors driving community variation.

Important findings Vegetation cover was significantly lower in swamp wetlands than in riverine wetlands, whereas soil electrical conductivity, total carbon, and total nitrogen contents were significantly higher. In total, 2 799 amplicon sequence variants (ASVs) belonging to the phylum Glomeromycota were identified and classified into 9 families and 10 genera, with Claroideoglomus and Glomus being the main predominant taxa. AMF α-diversity did not differ significantly between wetland types, whereas β-diversity showed significant differences. Variance partitioning analysis (VPA) indicated that wetland type and abiotic factors jointly explained 4.67% of community variation, with abiotic factors alone accounting for 3.39%, which was higher than the independent effect of wetland type (1.04%). Likewise, plant species and abiotic factors together explained 5.00% of AMF community variation, with the independent contribution of abiotic factors (2.04%) being slightly higher than that of plant species (1.38%). Mantel tests and canonical correspondence analysis (CCA) further demonstrated that plant richness, soil moisture, soil nutrient contents, and elevation were the key drivers of AMF community structure. In summary, AMF community structure in the alpine wetlands of the Yellow River Source Region is primarily driven by abiotic factors, while also being synergistically influenced by biotic components. These findings provide important scientific support for the development of targeted wetland ecological management strategies.

Key words: alpine wetland, arbuscular mycorrhizal fungi, community diversity, driving factor

ZHOU Chun-Han, XIONG Zhi-Cheng, YANG Ming-Xin, SHI Hai-Lan, ZHOU Ya-Xing, TANG Yu, ZHANG Jing, JI Bao-Ming, DAI Xin-Ling. Community characteristics and drivers of arbuscular mycorrhizal fungi in alpine wetlands of the Yellow River Source Region[J]. Chin J Plant Ecol, 2026, 50(3): 625-638.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2025.0342

https://www.plant-ecology.com/EN/Y2026/V50/I3/625

Figures/Tables 7

Fig. 1 Distribution map of sampling site in alpine wetlands of the Yellow River Source Region.

Table 1 Information of the sampling sites in alpine wetlands of the Yellow River Source Region

样点 Sampling site	经度 Longitude (E)	纬度 Latitude (N)	海拔 Altitude (m)	优势植物 Dominant species	湿地类型 Wetland type
S01	98.01°	35.07°	4 248	无脉薹草 Carex enervis	沼泽湿地 Swamp wetland
S02	98.17°	34.66°	4 212	无脉薹草 Carex enervis	河流湿地 Riverine wetland
S03	98.55°	34.37°	4 180	无脉薹草 Carex enervis	沼泽湿地 Swamp wetland
S04	97.92°	35.09°	4 255	无脉薹草 Carex enervis	沼泽湿地 Swamp wetland
S05	97.05°	34.83°	4 367	西藏嵩草 Carex tibetikobresia	沼泽湿地 Swamp wetland
S06	98.41°	34.17°	4 333	西藏嵩草 Carex tibetikobresia	河流湿地 Riverine wetland
S07	98.78°	35.26°	4 093	尖苞薹草 Carex microglochin	沼泽湿地 Swamp wetland
S08	98.40°	35.25°	4 120	尖苞薹草 Carex microglochin	沼泽湿地 Swamp wetland
S09	98.11°	34.77°	4 245	高山嵩草 Carex parvula	沼泽湿地 Swamp wetland
S10	98.03°	34.64°	4 222	高山嵩草 Carex parvula	沼泽湿地 Swamp wetland
S11	97.59°	34.59°	4 322	线叶嵩草 Carex capillifolia	河流湿地 Riverine wetland
S12	97.50°	34.79°	4 272	华扁穗草 Blysmus sinocompressus	沼泽湿地 Swamp wetland

Table 2 Plant characteristics, soil physical and chemical properties, and arbuscular mycorrhizal fungi (AMF) in plots with different dominant species and wetland types (mean ± SE)

Fig. 2 Relative abundance of arbuscular mycorrhizal fungi (AMF) community in the alpine wetlands of the Yellow River Source Region at the genus level: classified by wetland type (A) and dominant plant species (B). Different lowercase letters indicate significant differences (p < 0.05) in the relative abundance of the same AMF genus among dominant species.

Fig. 3 Principal coordinates analysis (PCoA) and permutational multivariate analysis of variance (PERMANOVA) of arbuscular mycorrhizal fungi (AMF) community across different wetlands types (A) and vegetation types (B).

Fig. 4 Variance partitioning analysis (VPA) of arbuscular mycorrhizal fungi (AMF) community with different wetland types (A) and dominant species (B).

Fig. 5 Mantel test correlation of plant communities, soil physicochemical properties, and AMF communities (A). *, p < 0.05; **, p < 0.01; ***, p < 0.001. Canonical correspondence analysis (CCA) showing key environmental factors influencing AMF community structure (B). Red lines in B indicate significant correlations, while black lines indicate non-significant correlation. AN, available nitrogen content; AP, available phosphorus content; EC, electrical conductivity; pH, soil pH; SOC, soil organic carbon content; SWC, soil water content; TN, total nitrogen content; TP, total phosphorus content.

References 78

[1]	Ba L, Ning JX, Wang DL, Facelli E, Facelli JM, Yang YN, Zhang LC (2012). The relationship between the diversity of arbuscular mycorrhizal fungi and grazing in a meadow steppe. Plant and Soil, 352, 143-156. DOI URL
[2]	Bhadalung NN, Suwanarit A, Dell B, Nopamornbodi O, Thamchaipenet A, Rungchuang J (2005). Effects of long-term NP-fertilization on abundance and diversity of arbuscular mycorrhizal fungi under a maize cropping system. Plant and Soil, 270, 371-382. DOI URL
[3]	Boddington CL, Bassett EE, Jakobsen I, Dodd JC (1999). Comparison of techniques for the extraction and quantification of extraradical mycelium of arbuscular mycorrhizal fungi in soils. Soil Biology & Biochemistry, 31, 479-482.
[4]	Brundrett M, Melville L, Peterson L (1994). Practical Methods in Mycorrhiza Research. Mycologue Publications, Sidney, Canada.
[5]	Cai XB, Peng YL (2016). Geographical distribution of arbuscular mycorrhizal fungi in the grassland ecosystems of the Tibetan Plateau. Acta Ecologica Sinica, 36, 2807-2818.
	[蔡晓布, 彭岳林 (2016). 西藏高原草地生态系统丛枝菌根真菌的地理分布. 生态学报, 36, 2807-2818.]
[6]	Chen KY, Xing S, Shi HL, Tang Y, Yang MX, Gu Q, Li YM, Zhang J, Ji BM (2023). Long-term fencing can’t benefit plant and microbial network stability of alpine meadow and alpine steppe in Three-River-Source National Park. Science of the Total Environment, 902, 166076. DOI: 10.1016/j.scitotenv.2023.166076.
[7]	Chiocchio V, Venedikian N, Martínez AE, Menéndez A, Ocampo JA, Godeas A (2000). Effect of the fungicide benomyl on spore germination and hyphal length of the arbuscular mycorrhizal fungus Glomus mosseae. International Microbiology, 3, 173-175.
[8]	Choudhury B, Kalita MC, Azad P (2010). Distribution of arbuscular mycorrhizal fungi in marshy and shoreline vegetation of Deepar Beel Ramsar Site of Assam, India. World Journal of Microbiology & Biotechnology, 26, 1965-1971. DOI URL
[9]	Coban O, de Deyn GB, van der Ploeg M (2022). Soil microbiota as game-changers in restoration of degraded lands. Science, 375, abe0725. DOI: 10.1126/science.abe0725.
[10]	Cornwell WK, Bedford BL, Chapin CT (2001). Occurrence of arbuscular mycorrhizal fungi in a phosphorus-poor wetland and mycorrhizal response to phosphorus fertilization. American Journal of Botany, 88, 1824-1829. PMID
[11]	Dashtebani F, Hajiboland R, Aliasgharzad N (2014). Characterization of salt-tolerance mechanisms in mycorrhizal (Claroideoglomus etunicatum) halophytic grass, Puccinellia distans. Puccinellia distans. Acta Physiologiae Plantarum, 36, 1713-1726.
[12]	Dolezal J, Dvorsky M, Kopecky M, Liancourt P, Hiiesalu I, Macek M, Altman J, Chlumska Z, Rehakova K, Capkova K, Borovec J, Mudrak O, Wild J, Schweingruber F (2016). Vegetation dynamics at the upper elevational limit of vascular plants in Himalaya. Scientific Reports, 6, 24881. DOI: 10.1038/srep24881.
[13]	Dolinar N, Regvar M, Abram D, Gaberščik A (2016). Water-level fluctuations as a driver of Phragmites australis primary productivity, litter decomposition, and fungal root colonisation in an intermittent wetland. Hydrobiologia, 774, 69-80. DOI URL
[14]	Domeignoz-Horta LA, Cappelli SL, Shrestha R, Gerin S, Lohila AK, Heinonsalo J, Nelson DB, Kahmen A, Duan PP, Sebag D, Verrecchia E, Laine AL (2024). Plant diversity drives positive microbial associations in the rhizosphere enhancing carbon use efficiency in agricultural soils. Nature Communications, 15, 8065. DOI: 10.1038/s41467-024-52449-5.
[15]	Du L, Liu SZ, Wang SQ, Yang J, Tang B, Li K, Zhou JC, Xu Y, Ye YF, Yang YQ, Tang H (2025). Arbuscular mycorrhizal fungi-mediated formation of different carbon fractions in various wetland types enhances carbon sequestration post-restoration on the Tibetan Plateau. Catena, 252, 108847. DOI: 10.1016/j.catena.2025.108847.
[16]	Egan CP, Callaway RM, Hart MM, Pither J, Klironomos J (2017). Phylogenetic structure of arbuscular mycorrhizal fungal communities along an elevation gradient. Mycorrhiza, 27, 273-282. DOI PMID
[17]	Fuchs B, Haselwandter K (2004). Red list plants: colonization by arbuscular mycorrhizal fungi and dark septate endophytes. Mycorrhiza, 14, 277-281. PMID
[18]	Gaberščik A, Krek JL, Zelnik I (2018). Habitat diversity along a hydrological gradient in a complex wetland results in high plant species diversity. Ecological Engineering, 118, 84-92. DOI URL
[19]	Gai JP, Tian H, Yang FY, Christie P, Li XL, Klironomos JN (2012). Arbuscular mycorrhizal fungal diversity along a Tibetan elevation gradient. Pedobiologia, 55, 145-151. DOI URL
[20]	Guo XJ, Shao QQ, Yang F, Li YZ, Wang YC, Wang DL (2019). Using UAV remote sensing for a population census of blue sheep (Pseudois nayaur) in Maduo County, source region of the Yellow River. Journal of Natural Resources, 34, 1054-1065. DOI URL
	[郭兴健, 邵全琴, 杨帆, 李愈哲, 汪阳春, 王东亮 (2019). 无人机遥感调查黄河源玛多县岩羊数量及分布. 自然资源学报, 34, 1054-1065.] DOI
[21]	Hart M, Reader R (2002). Host plant benefit from association with arbuscular mycorrhizal fungi: variation due to differences in size of Mycelium. Biology and Fertility of Soils, 36, 357-366. DOI URL
[22]	Hu D, Jiang XQ, Dai ZC, Chen DY, Zhang Y, Qi SS, Du DL (2024). Arbuscular mycorrhizal fungi enhance the capacity of invasive Sphagneticola trilobata to tolerate herbicides. Chinese Journal of Plant Ecology, 48, 651-659. DOI URL
	[胡蝶, 蒋欣琪, 戴志聪, 陈戴一, 张雨, 祁珊珊, 杜道林 (2024). 丛枝菌根真菌提高入侵杂草南美蟛蜞菊对除草剂的耐受性. 植物生态学报, 48, 651-659.] DOI
[23]	Jakobsen I, Abbott LK, Robson AD (1992). External hyphae of vesicular-arbuscular mycorrhizal fungi associated with Trifolium subterraneum L.: 2. Hyphal transport of ³²P over defined distances. New Phytologist, 120, 509-516. DOI URL
[24]	Jiang SJ, Liu YJ, Luo JJ, Qin MS, Johnson NC, Öpik M, Vasar M, Chai YX, Zhou XL, Mao L, Du GZ, An LZ, Feng HY (2018). Dynamics of arbuscular mycorrhizal fungal community structure and functioning along a nitrogen enrichment gradient in an Alpine meadow ecosystem. New Phytologist, 220, 1222-1235. DOI PMID
[25]	Johnson D, Vandenkoornhuyse PJ, Leake JR, Gilbert L, Booth RE, Grime JP, Young JPW, Read DJ (2004). Plant communities affect arbuscular mycorrhizal fungal diversity and community composition in grassland microcosms. New Phytologist, 161, 503-515. DOI PMID
[26]	Johnson NC (2010). Resource stoichiometry elucidates the structure and function of arbuscular mycorrhizas across scales. New Phytologist, 185, 631-647. DOI PMID
[27]	Johnson NC, Wolf J, Koch GW (2003). Interactions among mycorrhizae, atmospheric CO₂ and soil N impact plant community composition. Ecology Letters, 6, 532-540. DOI URL
[28]	Kang LY, Song YT, Mackelprang R, Zhang DY, Qin SQ, Chen LY, Wu LW, Peng YF, Yang YH (2024). Metagenomic insights into microbial community structure and metabolism in alpine permafrost on the Tibetan Plateau. Nature Communications, 15, 5920. DOI: 10.1038/S41467-024-50276-2.
[29]	Li CC, Zhou YX, Gu Q, Yang MX, Zhu CL, Peng ZY, Xue K, Zhao XQ, Wang YF, Ji BM, Zhang J (2021). The species diversity and community assembly of arbuscular mycorrhizal fungi in typical alpine grassland in Sanjiangyuan region. Acta Prataculturae Sinica, 30(1), 46-58.
	[李聪聪, 周亚星, 谷强, 杨明新, 朱传鲁, 彭子原, 薛凯, 赵新全, 王艳芬, 纪宝明, 张静 (2021). 三江源区典型高寒草地丛枝菌根真菌多样性及构建机制. 草业学报, 30(1), 46-58.] DOI
[30]	Li T, Chen BD (2012). Arbuscular mycorrhizal fungi improving drought tolerance of maize plants by up-regulation of aquaporin gene expressions in roots and the fungi themselves. Chinese Journal of Plant Ecology, 36, 973-981. DOI
	[李涛, 陈保冬 (2012). 丛枝菌根真菌通过上调根系及自身水孔蛋白基因表达提高玉米抗旱性. 植物生态学报, 36, 973-981.]
[31]	Li XL, Zhang JL, Gai JP, Cai XB, Christie P, Li XL (2015). Contribution of arbuscular mycorrhizal fungi of sedges to soil aggregation along an altitudinal alpine grassland gradient on the Tibetan Plateau. Environmental Microbiology, 17, 2841-2857. DOI PMID
[32]	Liu S, Che LN, Wan LH, Zhang WX, Chen J (2025). Wetland types and soil properties shape microbial communities in permafrost-degraded swamps. Catena, 249, 108666. DOI: 10.1016/j.catena.2024.108666.
[33]	Liu Y, Liu GH, Xiong ZQ, Liu WZ (2017). Response of greenhouse gas emissions from three types of wetland soils to simulated temperature change on the Qinghai-Tibetan Plateau. Atmospheric Environment, 171, 17-24. DOI URL
[34]	Liu Y, Shi AQ, Chen Y, Xu ZH, Liu YX, Yao YL, Wang YM, Jia BL (2024). Beneficial microorganisms: regulating growth and defense for plant welfare. Plant Biotechnology Journal, 23, 986-998. DOI URL
[35]	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: diverse AM fungi in Tibet Plateau. FEMS Microbiology Ecology, 78, 355-365. DOI PMID
[36]	Liu YJ, Shi GX, Mao L, Cheng G, Jiang SJ, Ma XJ, An LZ, Du GZ, Collins Johnson N, 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
[37]	Lu RK (2000). Soil Agricultural Chemical Analysis Methods. China Agricultural Science and Technology Press, Beijing. 12-14.
	[鲁如坤 (2000). 土壤农业化学分析方法. 中国农业科技出版社, 北京. 12-14.]
[38]	Ma YT, Li JQ, Wu J, Kong ZY, Feinstein LM, Ding X, Ge G, Wu L (2018). Bacterial and fungal community composition and functional activity associated with lake wetland water level gradients. Scientific Reports, 8, 760. DOI: 10.1038/s41598-018-19153-z. PMID
[39]	Mao H, Whalen JK, Zhang ZK, Sheng XJ, Hu GR, Chen B, Ma MJ (2024). Arbuscular mycorrhizal fungi associated with alpine meadow multifunctionality in a warmer climate with variable precipitation. Soil Biology & Biochemistry, 198, 109555. DOI: 10.1016/j.soilbio.2024.109555.
[40]	Mayerhofer MS, Kernaghan G, Harper KA (2013). The effects of fungal root endophytes on plant growth: a meta-analysis. Mycorrhiza, 23, 119-128. DOI PMID
[41]	Miller SP (2000). Arbuscular mycorrhizal colonization of semi-aquatic grasses along a wide hydrologic gradient. New Phytologist, 145, 145-155. DOI URL
[42]	Miller SP, Sharitz RR (2000). Manipulation of flooding and arbuscular mycorrhiza formation influences growth and nutrition of two semiaquatic grass species. Functional Ecology, 14, 738-748. DOI URL
[43]	Nelson DW (1996). Total carbon, organic carbon, and organic matter//Sparks DL. Methods of Soil Analysis, Part 3: Chemical Methods. Soil Science Society of America and American Society of Agronomy, Madison, USA.
[44]	Niu ZG, Jing YH, Zhang DQ, Zhang B (2024). An overview and the outlook for wetland ecosystems in the Qinghai-Tibetan Plateau under climate change. Climate Change Research, 20, 509-518.
	[牛振国, 景雨航, 张东启, 张波 (2024). 气候变化背景下青藏高原湿地生态系统响应特征: 回顾与展望. 气候变化研究进展, 20, 509-518.]
[45]	Niu ZG, Zhang HY, Wang XW, Yao WB, Zhou DM, Zhao KY, Zhao H, Li NN, Huang HB, Li CC, Yang J, Liu CX, Liu S, Wang L, Li Z, et al. (2012). Mapping wetland changes in China between 1978 and 2008. Chinese Science Bulletin, 57, 2813-2823. DOI URL
[46]	Ohsowski BM, Zaitsoff PD, Öpik M, Hart MM (2014). Where the wild things are: looking for uncultured Glomeromycota. New Phytologist, 204, 171-179. DOI PMID
[47]	Oliveira RS, Dodd JC, Castro PML (2001). The mycorrhizal status of Phragmites australis in several polluted soils and sediments of an industrialised region of Northern Portugal. Mycorrhiza, 10, 241-247.
[48]	Ouyang JX, Li W, Cao Y (2024). Preliminary study on factors driving arbuscular mycorrhizal fungi diversity with elevation in the South-Xizang River Basin. Plant Science Journal, 42(1), 34-42.
	[欧阳建鑫, 李伟, 操瑜 (2024). 藏南谷地湿地植物丛枝菌根真菌群落结构随海拔变化的驱动因素初探. 植物科学学报, 42(1), 34-42.]
[49]	Peng YL, Cai XB (2015). Changes of arbuscular mycorrhizal fungal community in an alpine grassland altitudinal gradient. Acta Ecologica Sinica, 35, 7475-7484.
	[彭岳林, 蔡晓布 (2015). 丛枝菌根真菌群落沿高寒草原海拔梯度的变化特征. 生态学报, 35, 7475-7484.]
[50]	Saint-Etienne L, Paul S, Imbert D, Dulormne M, Muller F, Toribio A, Plenchette C, Bâ AM (2006). Arbuscular mycorrhizal soil infectivity in a stand of the wetland tree Pterocarpus officinalis along a salinity gradient. Forest Ecology and Management, 232, 86-89. DOI URL
[51]	Schenck NC, Smith GS (1982). Responses of six species of vesicular-arbuscular mycorrhizal fungi and their effects on soybean at four soil temperatures. New Phytologist, 92, 193-201. DOI URL
[52]	Shi GX, Yao BQ, Liu YJ, Pan JB, Jiang SJ, Wang YB, Wang ZB, Feng HY, Zhou HK (2021). The effects of long-term warming on arbuscular mycorrhizal fungal communities depend on habitat type on the Qinghai-Tibet Plateau. Applied Soil Ecology, 167, 104030. DOI: 10.1016/j.apsoil.2021.104030.
[53]	Silvani VA, Colombo RP, Scorza MV, Fernández Bidondo L, Rothen CP, Scotti A, Fracchia S, Godeas A (2017). Arbuscular mycorrhizal fungal diversity in high-altitude hypersaline Andean wetlands studied by 454-sequencing and morphological approaches. Symbiosis, 72, 143-152. DOI URL
[54]	Smith SE, Read DJ (2008). Mycorrhizal Symbiosis. 3rd ed. Academic Press, London.
[55]	Smith SE, Smith FA (2011). Roles of arbuscular mycorrhizas in plant nutrition and growth: new paradigms from cellular to ecosystem scales. Annual Review of Plant Biology, 62, 227-250. DOI PMID
[56]	Stephen DT, James PA, Robert MS, Carl FF (2000). Mycorrhizal fungi associated with plants in ground-water fed wetlands. Wetlands, 20, 200-204. DOI URL
[57]	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. DOI URL
[58]	Taylor J, Harrier L (2000). A comparison of nine species of arbuscular mycorrhizal fungi on the development and nutrition of micropropagated Rubus idaeus L. cv. Glen Prosen (Red Raspberry). Plant and Soil, 225, 53-61. DOI
[59]	Vallino M, Fiorilli V, Bonfante P (2014). Rice flooding negatively impacts root branching and arbuscular mycorrhizal colonization, but not fungal viability. Plant, Cell & Environment, 37, 557-572. DOI URL
[60]	van der Heijden MGA, Bardgett RD, van Straalen NM (2008). The unseen majority: soil microbes as drivers of plant diversity and productivity in terrestrial ecosystems. Ecology Letters, 11, 296-310. DOI PMID
[61]	van der Heijden MGA, Klironomos JN, Ursic M, Moutoglis P, Streitwolf-Engel R, Boller T, Wiemken A, Sanders IR (1998). Mycorrhizal fungal diversity determines plant biodiversity, ecosystem variability and productivity. Nature, 396, 69-72. DOI
[62]	van der Heyde MGA, Ohsowski B, Abbott LK, Hart M (2017). Arbuscular mycorrhizal fungus responses to disturbance are context-dependent. Mycorrhiza, 27, 431-440. DOI PMID
[63]	van Geel M, Jacquemyn H, Plue J, Saar L, Kasari L, Peeters G, van Acker K, Honnay O, Ceulemans T (2018). Abiotic rather than biotic filtering shapes the arbuscular mycorrhizal fungal communities of European seminatural grasslands. New Phytologist, 220, 1262-1272. DOI PMID
[64]	Wagg C, Jansa J, Schmid B, van der Heijden MGA (2011). Belowground biodiversity effects of plant symbionts support aboveground productivity: biodiversity effects of soil symbionts. Ecology Letters, 14, 1001-1009. DOI PMID
[65]	Wang YT, Qiu Q, Yang ZY, Hu ZJ, Tam NF, Xin GR (2010). Arbuscular mycorrhizal fungi in two mangroves in South China. Plant and Soil, 331, 181-191. DOI URL
[66]	Wei D, Xu-Ri, Tenzin-Tarchen, Wang YS, Wang YH (2015). Considerable methane uptake by alpine grasslands despite the cold climate: in situ measurements on the central Tibetan Plateau, 2008-2013. Global Change Biology, 21, 777-788. DOI PMID
[67]	Wu MY, Franz EH, Chen SL (2001). Oxygen fluxes and ammonia removal efficiencies in constructed treatment wetlands. Water Environment Research, 73, 661-666. PMID
[68]	Wu QS, Srivastava AK, Zou YN (2013). AMF-induced tolerance to drought stress in Citrus: a review. Scientia Horticulturae, 164, 77-87. DOI URL
[69]	Wu S, Sui X, Zhang T, Chen YT, Zhu DG, Cui FX, Yang LB (2019). Research on the progress of AMF in wetland. Territory & Natural Resources Study, (6), 80-84.
	[吴松, 隋心, 张童, 陈雨彤, 朱道光, 崔福星, 杨立宾 (2019). 湿地丛枝菌根真菌研究进展. 国土与自然资源研究, (6), 80-84.]
[70]	Yan JZ, Wu FS, Feng HY (2008). Review on the relationship between wetland plants and arbuscular mycorrhizal fungi (AMF). Acta Botanica Boreali-Occidentalia Sinica, 28, 4836-4842.
	[鄢金灼, 武发思, 冯虎元 (2008). 湿地植物与丛枝菌根真菌(AMF)相互关系的研究进展. 西北植物学报, 28, 4836-4842.]
[71]	Yang R, Qin ZF, Wang JJ, Xu S, Zhao W, Zhang XX, Huang ZY (2020). Salinity changes root occupancy by arbuscular mycorrhizal fungal species. Pedobiologia, 81, 150665. DOI: 10.1016/j.pedobi.2020.150665.
[72]	Zhang B, Li QY, Jing YH, Niu ZG, Gong P, Zhang DQ (2025). Alpine wetland distribution patterns and decreasing trends in the Qinghai-Tibetan Plateau. Science Bulletin, 70, 3509-3511. DOI URL
[73]	Zhang B, Xue K, Zhou ST, Che RX, Du JQ, Tang L, Pang Z, Wang F, Wang D, Cui XY, Hao YB, Wang YF (2019). Phosphorus mediates soil prokaryote distribution pattern along a small-scale elevation gradient in Noijin Kangsang Peak, Tibetan Plateau. FEMS Microbiology Ecology, 95, fiz076. DOI: 10.1093/femsec/fiz076.
[74]	Zhang J, Tang XL, Zhong SY, Yin GC, Gao YF, He XH (2017). Recalcitrant carbon components in glomalin-related soil protein facilitate soil organic carbon preservation in tropical forests. Scientific Reports, 7, 2391. DOI: 10.1038/s41598-017-02486-6.
[75]	Zhang Q, Sun QX, Koide RT, Peng ZH, Zhou JX, Gu XG, Gao WD, Yu M (2014). Arbuscular mycorrhizal fungal mediation of plant-plant interactions in a marshland plant community. The Scientific World Journal, 2014, 923610. DOI: 10.1155/2014/923610.
[76]	Zhao L, Zou DF, Hu GJ, Du EJ, Pang QQ, Xiao Y, Li R, Sheng Y, Wu XD, Sun Z, Wang LX, Wang C, Ma L, Zhou HY, Liu SB (2020). Changing climate and the permafrost environment on the Qinghai-Tibet (Xizang) Plateau. Permafrost and Periglacial Processes, 31, 396-405. DOI
[77]	Zheng J, Dong DH, Dong X, et al. (2015). China Wetland Resources: Qinghai Volume. China Forestry Publishing House, Beijing.
	[郑杰, 董得红, 董旭 (2015). 中国湿地资源·青海卷. 中国林业出版社, 北京.]
[78]	Zhou YX, Chen KY, Muneer MA, Li CC, Shi HL, Tang Y, Zhang J, Ji BM (2022). Soil moisture and pH differentially drive arbuscular mycorrhizal fungal composition in the riparian zone along an alpine river of Nam Co watershed. Frontiers in Microbiology, 13, 994918. DOI: 10.3389/fmicb.2022.994918.