, Volume 191, Issue 3, pp 697–708 | Cite as

Asymmetric responses of plant community structure and composition to precipitation variabilities in a semi-arid steppe

  • Mingxing Zhong
  • Jian Song
  • Zhenxing Zhou
  • Jingyi Ru
  • Mengmei Zheng
  • Ying Li
  • Dafeng Hui
  • Shiqiang WanEmail author
Global change ecology – original research


Changing precipitation regimes can profoundly affect plant growth in terrestrial ecosystems, especially in arid and semi-arid regions. However, how changing precipitation, especially extreme precipitation events, alters plant diversity and community composition is still poorly understood. A 3-year field manipulative experiment with seven precipitation treatments, including − 60%, − 40%, − 20%, 0% (as a control), + 20%, + 40%, and + 60% of ambient growing-season precipitation, was conducted in a semi-arid steppe in the Mongolian Plateau. Results showed total plant community cover and forb cover were enhanced with increased precipitation and reduced under decreased precipitation, whereas grass cover was suppressed under the − 60% treatment only. Plant community and grass species richness were reduced by the − 60% treatment only. Moreover, our results demonstrated that total plant community cover was more sensitive to decreased than increased precipitation under normal and extreme precipitation change, and species richness was more sensitive to decreased than increased precipitation under extreme precipitation change. The community composition and low field water holding capacity may drive this asymmetric response. Accumulated changes in community cover may eventually lead to changes in species richness. However, compared to control, Shannon–Weiner index (H) did not respond to any precipitation treatment, and Pielou’s evenness index (E) was reduced under the + 60% treatment across the 3 year, but not in each year. Thus, the findings suggest that plant biodiversity in the semi-arid steppe may have a strong resistance to precipitation pattern changes through adjusting its composition in a short term.


Dominated species Extreme precipitation Functional groups Sensitivity Species richness 



The authors thank Qian Zhang, Hongyan Han, and Fanglong Su for their help in the field experiments.

Author contribution statement

MXZ and SW conceived the ideas and designed methodology; MXZ, YM, AZ, ZZ, JR, MMZ, YL collected the data; MXZ and SW analyzed the data; MXZ, DH, and SW led the writing of the manuscript. All authors contributed critically to the drafts and gave final approval for publication.


This study was funded by the National Natural Science Foundation of China (31430015 and 31830012).

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Supplementary material

442_2019_4520_MOESM1_ESM.docx (194 kb)
Supplementary material 1 (DOCX 194 kb)


  1. Adler PB, Levine JM (2007) Contrasting relationships between precipitation and species richness in space and time. Oikos 116:221–232. CrossRefGoogle Scholar
  2. Báez S, Collins SL, Pockman WT, Johnson JE, Small EE (2012) Effects of experimental rainfall manipulations on Chihuahuan desert grassland and shrubland plant communities. Oecologia 172:1117–1127. CrossRefPubMedGoogle Scholar
  3. Bai YF, Wu JG, Xing Q, Pan QM, Huang JH, Yang DL, Han XG (2008) Primary production and rain use efficiency across a precipitation gradient on the Mongolia plateau. Ecology 89:2140–2153. CrossRefPubMedGoogle Scholar
  4. Bodner GS, Robles MD (2017) Enduring a decade of drought: patterns and drivers of vegetation change in a semi-arid grassland. J Arid Environ 136:1–14. CrossRefGoogle Scholar
  5. Bollig C, Feller U (2014) Impacts of drought stress on water relations and carbon assimilation in grassland species at different altitudes. Agric Ecosyst Environ 188:212–220. CrossRefGoogle Scholar
  6. Chen H, Ma L, Xin X, Liu J, Wang R (2018) Plant community responses to increased precipitation and belowground litter addition: evidence from a 5-year semiarid grassland experiment. Ecol Evol 8:4587–4597. CrossRefPubMedPubMedCentralGoogle Scholar
  7. Cleland EE, Chiariello NR, Loarie SR, Mooney HA, Field CB (2006) Diverse responses of phenology to global changes in a grassland ecosystem. Proc Natl Acad Sci USA 103:13740–13744. CrossRefPubMedGoogle Scholar
  8. Copeland SM, Harrison SP, Latimer AM, Damschen EI, Eskelinen AM, Fernandez-Going B, Spasojevic MJ, Anacker BL, Thorne JH (2016) Ecological effects of extreme drought on Californian herbaceous plant communities. Ecol Monogr 86:295–311. CrossRefGoogle Scholar
  9. De Boeck HJ, Dreesen FE, Janssens IA, Nijs I (2011) Whole-system responses of experimental plant communities to climate extremes imposed in different seasons. New Phytol 189:806–817. CrossRefGoogle Scholar
  10. Deng Q, Aras S, Yu CL, Dzantor EK, Fay PA, Luo YQ, Shen WJ, Hui DF (2017) Effects of precipitation changes on aboveground net primary production and soil respiration in a switchgrass field. Agric Ecosyst Environ 248:29–37. CrossRefGoogle Scholar
  11. Epstein HE, Lauenroth WK, Burke IC (1997) Effects of temperature and soil texture on ANPP in the US Great Plains. Ecology 78:2628–2631CrossRefGoogle Scholar
  12. Fay PA, Carlisle JD, Knapp AK, Blair JM, Collins SL (2003) Productivity responses to altered rainfall patterns in a C4 -dominated grassland. Oecologia 137:245–251. CrossRefPubMedGoogle Scholar
  13. Felton AJ, Zavislan-Pullaro S, Smith MD (2019) Semiarid ecosystem sensitivity to precipitation extremes: weak evidence for vegetation constraints. Ecology 100:e02572. CrossRefPubMedGoogle Scholar
  14. Fu G, Shen Z, Zhang X (2018) Increased precipitation has stronger effects on plant production of an alpine meadow than does experimental warming in the Northern Tibetan Plateau. Agric For Meteorol 249:11–21. CrossRefGoogle Scholar
  15. Gherardi LA, Sala OE (2015) Enhanced precipitation variability decreases grass- and increases shrub-productivity. Proc Natl Acad Sci USA 112:12735–12740. CrossRefPubMedGoogle Scholar
  16. Hsu JS, Powell J, Adler PB (2012) Sensitivity of mean annual primary production to precipitation. Glob Chang Biol 18:2246–2255. CrossRefGoogle Scholar
  17. Hui D, Jackson RB (2006) Geographical and interannual variability in biomass partitioning in grassland ecosystems: a synthesis of field data. New Phytol 169:85–93. CrossRefPubMedGoogle Scholar
  18. Huxman TE, Smith MD, Fay PA, Knapp AK, Shaw MR, Loik ME, Smith SD, Tissue DT, Zak JC, Weltzin JF (2004) Convergence across biomes to a common rain-use efficiency. Nature 429:651–654. CrossRefPubMedGoogle Scholar
  19. IPCC (2013) The physical science basis. Contribution of working group I to the fifth assessment report of the intergovernmental panel on climate change. Cambridge University Press, CambridgeGoogle Scholar
  20. Jobbágy EG, Sala OE (2000) Controls of grass and shrub aboveground production in the patagonian steppe. Ecol Appl 10:541–549CrossRefGoogle Scholar
  21. Jost L (2006) Entropy and diversity. Oikos 113:363–375. CrossRefGoogle Scholar
  22. Knapp AK, Briggs JM, Koelliker JK (2001) Frequency and extent of water limitation to primary production in a Mesic temperate grassland. Ecosystems 4:19–28. CrossRefGoogle Scholar
  23. Knapp AK, Fay PA, Blair JM, Collins SL, Smith MD, Carlisle JD, Harper CW, Danner BT, Lett MS, McCarron JK (2002) Rainfall variability, carbon cycling, and plant species diversity in a mesic grassland. Science 298:2202–2205. CrossRefPubMedGoogle Scholar
  24. Knapp AK, Beier C, Briske DD, Classen AT, Luo YQ, Reichstein M, Smith MD, Smith SD, Bell JE, Fay PA (2008) Consequences of more extreme precipitation regimes for terrestrial ecosystems. Bioscience 58:811–821. CrossRefGoogle Scholar
  25. Knapp AK, Ciais P, Smith MD (2017) Reconciling inconsistencies in precipitation-productivity relationships: implications for climate change. New Phytol 214:41–47. CrossRefGoogle Scholar
  26. La Pierre KJ, Blumenthal DM, Brown CS, Klein JA, Smith MD (2016) Drivers of variation in aboveground net primary productivity and plant community composition differ across a broad precipitation gradient. Ecosystems 19:521–533. CrossRefGoogle Scholar
  27. Ladwig LM, Collins SL, Swann AL, Xia Y, Allen MF, Allen EB (2012) Above- and belowground responses to nitrogen addition in a Chihuahuan desert grassland. Oecologia 169:177–185. CrossRefPubMedGoogle Scholar
  28. Liu H, Mi Z, Lin L, Wang Y, Zhang Z, Zhang F, Wang H, Liu L, Zhu B, Cao G, Zhao X, Sanders NJ, Classen AT, Reich PB, He JS (2018) Shifting plant species composition in response to climate change stabilizes grassland primary production. Proc Natl Acad Sci USA 115:4051–4056. CrossRefPubMedGoogle Scholar
  29. Lloret F, Penuelas J, Estiarte M (2004) Experimental evidence of reduced diversity of seedlings due to climate modification in a Mediterranean-type community. Glob Chang Biol 10:248–258. CrossRefGoogle Scholar
  30. Luo Y, Jiang L, Niu S, Zhou X (2017) Nonlinear responses of land ecosystems to variation in precipitation. New Phytol 214:5–7. CrossRefPubMedGoogle Scholar
  31. Miao Y, Han H, Du Y, Zhang Q, Jiang L, Hui D, Wan S (2017) Nonlinear responses of soil respiration to precipitation changes in a semiarid temperate steppe. Sci Rep 7:45782. CrossRefPubMedPubMedCentralGoogle Scholar
  33. Robertson TR, Bell CW, Zak JC, Tissue DT (2009) Precipitation timing and magnitude differentially affect aboveground annual net primary productivity in three perennial species in a Chihuahuan Desert grassland. New Phytol 181:230–242. CrossRefPubMedGoogle Scholar
  34. Sala OE, Lauenroth WK, Golluscio RA (1997) Plant functional types in temperate semi-arid regions. In: Smith TM, Shugart HH, Woodward FI (eds) Plant functional types. Cambridge University Press, Cambridge, pp 217–233Google Scholar
  35. Schwinning S, Sala OE (2004) Hierarchy of responses to resource pulses in arid and semi-arid ecosystems. Oecologia 141:211–220. CrossRefPubMedGoogle Scholar
  36. Schwinning S, Starr BI, Ehleringer JR (2005) Summer and winter drought in a cold desert ecosystem (Colorado Plateau) part II: effects on plant carbon assimilation and growth. J Arid Environ 61:61–78. CrossRefGoogle Scholar
  37. Sherry RA, Arnone JA, Johnson DW, Schimel DS, Verburg PS, Luo Y (2011) Carry over from previous year environmental conditions alters dominance hierarchy in a prairie plant community. J Plant Ecol 5:134–146. CrossRefGoogle Scholar
  38. Shi Z, Lin Y, Wilcox KR, Souza L, Jiang L, Jiang J, Jung CG, Xu X, Yuan M, Guo X, Wu L, Zhou J, Luo Y (2018) Successional change in species composition alters climate sensitivity of grassland productivity. Glob Chang Biol 24:4993–5003. CrossRefPubMedGoogle Scholar
  39. Smith MD, Knapp AK (2003) Dominant species maintain ecosystem function with non-random species loss. Ecol Lett 6:509–517. CrossRefGoogle Scholar
  40. Smith MD, Wilcox KR, Power SAD, Tissue T, Knapp AK (2017) Assessing community and ecosystem sensitivity to climate change–toward a more comparative approach. J Veg Sci 28:235–237. CrossRefGoogle Scholar
  41. Solomon S, Plattner G-K, Knutti R, Friedlingstein P (2009) Irreversible climate change due to carbon dioxide emissions. Proc Natl Acad Sci USA 106:1704–1709CrossRefGoogle Scholar
  42. Storch D, Bohdalkova E, Okie J (2018) The more-individuals hypothesis revisited: the role of community abundance in species richness regulation and the productivity-diversity relationship. Ecol Lett 21:920–937. CrossRefPubMedGoogle Scholar
  43. Wilcox KR, von Fischer JC, Muscha JM, Petersen MK, Knapp AK (2015) Contrasting above- and belowground sensitivity of three Great Plains grasslands to altered rainfall regimes. Glob Chang Biol 21:335–344. CrossRefPubMedGoogle Scholar
  44. Wilcox KR, Shi Z, Gherardi LA, Lemoine NP, Koerner SE, Hoover DL, Bork E, Byrne KM, Cahill JJ, Collins SL et al (2017) Asymmetric responses of primary productivity to precipitation extremes: a synthesis of grassland precipitation manipulation experiments. Glob Chang Biol 23:4376–4385. CrossRefPubMedGoogle Scholar
  45. Wu Z, Dijkstra P, Koch GW, PeÑUelas J, Hungate BA (2011) Responses of terrestrial ecosystems to temperature and precipitation change: a meta-analysis of experimental manipulation. Glob Chang Biol 17:927–942. CrossRefGoogle Scholar
  46. Wu D, Ciais P, Viovy N, Knapp AK, Wilcox K, Bahn M, Smith MD, Vicca S, Fatichi S, Zscheischler J et al (2018a) Asymmetric responses of primary productivity to altered precipitation simulated by ecosystem models across three long-term grassland sites. Biogeosciences 15:3421–3437. CrossRefGoogle Scholar
  47. Wu X, Liu H, Li X, Ciais P, Babst F, Guo W, Zhang C, Magliulo V, Pavelka M, Liu S, Huang Y, Wang P, Shi C, Ma Y (2018b) Differentiating drought legacy effects on vegetation growth over the temperate Northern Hemisphere. Glob Chang Biol 24:504–516. CrossRefPubMedGoogle Scholar
  48. Xia Y, Moore DI, Collins SL, Muldavin EH (2010) Aboveground production and species richness of annuals in Chihuahuan Desert grassland and shrubland plant communities. J Arid Environ 74:378–385. CrossRefGoogle Scholar
  49. Xu Z, Wan S, Ren H, Han X, Li MH, Cheng W, Jiang Y (2012) Effects of water and nitrogen addition on species turnover in temperate grasslands in northern China. PLoS One 7:e39762. CrossRefPubMedPubMedCentralGoogle Scholar
  50. Yahdjian L, Sala OE (2002) A rainout shelter design for intercepting different amounts of rainfall. Oecologia 133:95–101. CrossRefPubMedGoogle Scholar
  51. Yang H, Li Y, Wu M, Zhang Z, Li L, Wan S (2011a) Plant community responses to nitrogen addition and increased precipitation: the importance of water availability and species traits. Glob Chang Biol 17:2936–2944. CrossRefGoogle Scholar
  52. Yang H, Wu M, Liu W, Zhang Z, Zhang N, Wan S (2011b) Community structure and composition in response to climate change in a temperate steppe. Glob Chang Biol 17:452–465. CrossRefGoogle Scholar
  53. Yang H, Jiang L, Li L, Li A, Wu M, Wan S (2012) Diversity-dependent stability under mowing and nutrient addition: evidence from a 7-year grassland experiment. Ecol Lett 15:619–626. CrossRefPubMedGoogle Scholar
  54. Yang X, Yang ZL, Tan JQ, Li GY, Wan SQ, Jiang L, Shefferson R (2018) Nitrogen fertilization, not water addition, alters plant phylogenetic community structure in a semi-arid steppe. J Ecol 106:991–1000. CrossRefGoogle Scholar
  55. Ye JS, Pei JY, Fang C (2018) Under which climate and soil conditions the plant productivity-precipitation relationship is linear or nonlinear? Sci Total Environ 616–617:1174–1180. CrossRefPubMedGoogle Scholar
  56. Zeppel MJ, Adams HD, Anderegg WR (2011) Mechanistic causes of tree drought mortality: recent results, unresolved questions and future research needs. New Phytol 192:800–803. CrossRefPubMedGoogle Scholar
  57. Zeppel MJB, Wilks JV, Lewis JD (2014) Impacts of extreme precipitation and seasonal changes in precipitation on plants. Biogeosciences 11:3083–3093. CrossRefGoogle Scholar
  58. Zhang B, Zhu JJ, Pan QM, Liu YS, Chen SP, Chen DM, Yan Y, Dou SD, Han XG (2017a) Grassland species respond differently to altered precipitation amount and pattern. Environ Exp Bot 137:166–176. CrossRefGoogle Scholar
  59. Zhang FY, Quan Q, Song B, Sun J, Chen YJ, Zhou QP, Niu SL (2017b) Net primary productivity and its partitioning in response to precipitation gradient in an alpine meadow. Sci Rep 7:15193. CrossRefPubMedPubMedCentralGoogle Scholar
  60. Zhang L, Xie Z, Zhao R, Zhang Y (2018) Plant, microbial community and soil property responses to an experimental precipitation gradient in a desert grassland. Appl Soil Ecol 127:87–95. CrossRefGoogle Scholar
  61. Zhao C, Yuan M, Yu CD, Zhu LL, Wang F, Jiang L, Hui DF, Wan SQ (2016) Soil microbial community composition and respiration along an experimental precipitation gradient in a semiarid steppe. Sci Rep 6:24317. CrossRefPubMedPubMedCentralGoogle Scholar

Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2019

Authors and Affiliations

  1. 1.International Joint Research Laboratory for Global Change Ecology, College of Life SciencesHenan UniversityKaifengChina
  2. 2.College of Life ScienceHebei UniversityBaodingChina
  3. 3.Department of Biological SciencesTennessee State UniversityNashvilleUSA

Personalised recommendations