Decomposition time, chemical traits and climatic factors determine litter-mixing effects on decomposition in an alpine steppe ecosystem in Northern Tibet

  • Youchao Chen
  • Shuqin Ma
  • Hongmao Jiang
  • Degyi Yangzom
  • Genwei Cheng
  • Xuyang LuEmail author
Regular Article



In the context of rapid species diversity loss, whether and how species diversity affects litter decomposition in alpine ecosystems remain unclear. Here, we aimed to assess the impact of litter mixing on decomposition and the influencing factors of such litter-mixing effects (LMEs).


We incubated the litter of four alpine steppe species in monocultures and all possible litter combinations and analysed the mass remaining over three years.


Litter mixing showed more frequent antagonistic effects than synergistic effects in the early stage of the experiment; however, the LME might change from antagonistic to synergistic at the latter stage of incubation. The magnitude of the LMEs increased as decomposition progressed. The four-species mixture resulted in more antagonistic effects, while the magnitude of the non-additive effects showed no relationship with species richness. Litter chemical traits and climatic factors affected the direction and magnitude of the non-additive effects, but these influences were time-dependent. Additionally, the fast-decomposing and slow-decomposing litter species had negative and positive responses to the litter mixture, respectively.


Our results demonstrate that the LME was determined by decomposition time, chemical traits and climatic factors and suggest that assessing the relative individual performances of the litter components in a mixture is an important way to examine the mechanisms underlying LMEs.


Biodiversity Litter decomposition Litter mixture Northern Tibet Non-additive effects 



This study was supported by the Natural Science Foundation of China (Grant No. 41671262 &41877338), Key Laboratory of Mountain Surface Processes and Ecological Regulation, Chinese Academy of Sciences, and the Natural Science Foundation of Tibet Autonomous Region.

Supplementary material

11104_2019_4131_MOESM1_ESM.docx (12.6 mb)
ESM 1 (DOCX 12863 kb)


  1. Ball BA, Hunter MD, Kominoski JS et al (2014) Consequences of non-random species loss for decomposition dynamics: experimental evidence for additive and non-additive effects. J Ecol 96:303–313CrossRefGoogle Scholar
  2. Barantal S, Roy J, Fromin N, Schimann H, Hättenschwiler S (2011) Long-term presence of tree species but not chemical diversity affect litter mixture effects on decomposition in a neotropical rainforest. Oecologia 167:241–252CrossRefGoogle Scholar
  3. Butenschoen O, Krashevska V, Maraun M, Marian F, Sandmann D, Scheu S (2014) Litter mixture effects on decomposition in tropical montane rainforests vary strongly with time and turn negative at later stages of decay. Soil Biol Biochem 77:121–128CrossRefGoogle Scholar
  4. Cardinale BJ, Matulich KL, Hooper DU, Byrnes JE, Duffy E, Gamfeldt L, Balvanera P, O'Connor MI, Gonzalez A (2011) The functional role of producer diversity in ecosystems. Am J Bot 98:572–592CrossRefGoogle Scholar
  5. Chen Y, Sun J, Xie F, Wang X, Cheng G, Lu X (2015) Litter chemical structure is more important than species richness in affecting soil carbon and nitrogen dynamics including gas emissions from an alpine soil. Biol Fertil Soils 51:791–800CrossRefGoogle Scholar
  6. de Magalhães RMQ, Schwilk DW (2012) Leaf traits and litter flammability: evidence for non-additive mixture effects in a temperate forest. J Ecol 100:1153–1163CrossRefGoogle Scholar
  7. Duan J, Wang S, Zhang Z, Xu G, Luo C, Chang X, Zhu X, Cui S, Zhao X, Wang W, du M (2013) Non-additive effect of species diversity and temperature sensitivity of mixed litter decomposition in the alpine meadow on Tibetan plateau. Soil Biol Biochem 57:841–847CrossRefGoogle Scholar
  8. Foster P (2001) The potential negative impacts of global climate change on tropical montane cloud forests. Earth-Sci Rev 55:73–106CrossRefGoogle Scholar
  9. Frainer A, Moretti MS, Xu W, Gessner MO (2015) No evidence for leaf-trait dissimilarity effects on litter decomposition fungal decomposers and nutrient dynamics. Ecology 96:550–561CrossRefGoogle Scholar
  10. Gartner TB, Cardon ZG (2004) Decomposition dynamics in mixed-species leaf litter. Oikos 104:230–246CrossRefGoogle Scholar
  11. Gessner MO, Swan CM, Dang CK, McKie BG, Bardgett RD, Wall DH, Hättenschwiler S (2010) Diversity meets decomposition. Trends Ecol Evol 25:372–380CrossRefGoogle Scholar
  12. Hättenschwiler S, Tiunov AV, Scheu S (2005) Biodiversity and litter decomposition in terrestrial ecosystems. Annu Rev Ecol Evol Syst 36:191–218. CrossRefGoogle Scholar
  13. Hoorens B, Aerts R, Stroetenga M (2003) Does initial litter chemistry explain litter mixture effects on decomposition? Oecologia 137:578–586CrossRefGoogle Scholar
  14. Hong J, Wang X, Wu J (2014) Stoichiometry of root and leaf nitrogen and phosphorus in a dry alpine steppe on the northern Tibetan Plateau. PLoS One 9:e109052–8Google Scholar
  15. Joly FX, Milcu A, Scherer-Lorenzen M, Jean LK, Bussotti F, Dawud SM, Müller S, Pollastrini M, Raulund-Rasmussen K, Vesterdal L, Hättenschwiler S (2017) Tree species diversity affects decomposition through modified micro-environmental conditions across European forests. New Phytol 214:1281–1293CrossRefGoogle Scholar
  16. Keith AM, van der Wal R, Brooker RW, Osler GHR, Chapman SJ, Burslem DFRP, Elston DA (2008) Increasing litter species richness reduces variability. Ecology 89:2657–2665CrossRefGoogle Scholar
  17. Kominoski AJS, Pringle CM, B a B et al (2007) Nonadditive effects of leaf litter species diversity on breakdown dynamics in a detritus- based stream. Ecology 88:1167–1176CrossRefGoogle Scholar
  18. Lecerf A, Marie G, Kominoski JS, LeRoy CJ, Bernadet C, Swan CM (2011) Incubation time, functional litter diversity, and ecosystem characteristics predict nonadditive litter mixing effects on decomposition: a synthesis from streams. Ecology 92:160–169CrossRefGoogle Scholar
  19. Lin G, Zeng D (2018) Functional identity rather than functional diversity or species richness controls litter mixture decomposition in a subtropical forest. Plant Soil 428:179–193CrossRefGoogle Scholar
  20. Liu P, Sun OJ, Huang J, Li L, Han X (2007) Nonadditive effects of litter mixtures on decomposition and correlation with initial litter N and P concentrations in grassland plant species of northern China. Biol Fertil Soils 44:211–216CrossRefGoogle Scholar
  21. Liu C, Liu Y, Guo K, Zhao H, Qiao X, Wang S, Zhang L, Cai X (2016) Mixing litter from deciduous and evergreen trees enhances decomposition in a subtropical karst forest in southwestern China. Soil Biol Biochem 101:44–54CrossRefGoogle Scholar
  22. Makkonen M, Berg MP, van Logtestijn RSP, van Hal JR, Aerts R (2013) Do physical plant litter traits explain non-additivity in litter mixtures? A test of the improved microenvironmental conditions theory. Oikos 122:987–997CrossRefGoogle Scholar
  23. Mao B, Yu ZY, Zeng DH (2015) Non-additive effects of species mixing on litter mass loss and chemical properties in a Mongolian pine plantation of Northeast China. Plant Soil 396:339–351CrossRefGoogle Scholar
  24. Rosemond AD, Swan CM, Kominoski JS, Dye SE (2010) Non-additive effects of litter mixing are suppressed in a nutrient-enriched stream. Oikos 119:326–336CrossRefGoogle Scholar
  25. Santonja M, Baldy V, Fernandez C (2015a) Potential shift in plant communities with climate change : outcome on litter decomposition and nutrient release in a Mediterranean oak Forest. Ecosystems 18:1253–1268CrossRefGoogle Scholar
  26. Santonja M, Fernandez C, Gauquelin T, Baldy V (2015b) Climate change effects on litter decomposition: intensive drought leads to a strong decrease of litter mixture interactions. Plant Soil 393:69–82CrossRefGoogle Scholar
  27. Santonja M, Rancon A, Fromin N, Baldy V, Hättenschwiler S, Fernandez C, Montès N, Mirleau P (2017) Plant litter diversity increases microbial abundance, fungal diversity, and carbon and nitrogen cycling in a Mediterranean shrubland. Soil Biol Biochem 111:124–134CrossRefGoogle Scholar
  28. Schuster MJ, Dengler J, Kreyling J et al (2017) Drought inhibits synergistic interactions of native and exotic litter mixtures during decomposition in temperate grasslands. Plant Soil 415:257–268CrossRefGoogle Scholar
  29. Srivastava DS, Cardinale BJ, Downing AL, Duffy JE, Jouseau C, Sankaran M, Wright JP (2009) Diversity has stronger top-down than bottom-up effects on decomposition. Ecology 90:1073–1083CrossRefGoogle Scholar
  30. Thoss V, Shevtsova A, Nilsson MC (2004) Environmental manipulation treatment effects on the reactivity of water-soluble phenolics in a subalpine tundra ecosystem. Plant Soil 259:355–365CrossRefGoogle Scholar
  31. Tilman D, Isbell F, Cowles JM (2014) Biodiversity and ecosystem functioning. Annu Rev Ecol Evol Syst 45:471–493CrossRefGoogle Scholar
  32. Trogisch S, He JS, Hector A, Scherer-Lorenzen M (2016) Impact of species diversity, stand age and environmental factors on leaf litter decomposition in subtropical forests in China. Plant Soil 400:337–350CrossRefGoogle Scholar
  33. Vivanco L, Austin AT (2008) Tree species identity alters forest litter decomposition through long-term plant and soil interactions in Patagonia, Argentina. J Ecol 96:727–736CrossRefGoogle Scholar
  34. Wardle DA, Bonner KI, Nicholson KS (1997) Biodiversity and plant litter: experimental evidence which does not support the view that enhanced species richness improves ecosystem function. Oikos 79:247CrossRefGoogle Scholar
  35. Wu D, Li T, Wan S (2013) Time and litter species composition affect litter-mixing effects on decomposition rates. Plant Soil 371:355–366CrossRefGoogle Scholar
  36. Zeng L, He W, Teng M, Luo X, Yan Z, Huang Z, Zhou Z, Wang P, Xiao W (2018) Effects of mixed leaf litter from predominant afforestation tree species on decomposition rates in the three gorges reservoir, China. Sci Total Environ 639:679–686CrossRefGoogle Scholar
  37. Zhou G, Guan L, Wei X, Tang X, Liu S, Liu J, Zhang D, Yan J (2008) Factors influencing leaf litter decomposition: an intersite decomposition experiment across China. Plant Soil 311:61–72CrossRefGoogle Scholar

Copyright information

© Springer Nature Switzerland AG 2019

Authors and Affiliations

  • Youchao Chen
    • 1
    • 2
  • Shuqin Ma
    • 3
  • Hongmao Jiang
    • 2
    • 4
  • Degyi Yangzom
    • 5
  • Genwei Cheng
    • 2
  • Xuyang Lu
    • 2
    Email author
  1. 1.Key Laboratory of Aquatic Botany and Watershed Ecology, Wuhan Botanical GardenChinese Academy of SciencesWuhanChina
  2. 2.Institute of Mountain Hazards and EnvironmentChinese Academy of SciencesChengduChina
  3. 3.College of TourismHenan Normal UniversityXinxiangChina
  4. 4.University of Chinese Academy of SciencesBeijingChina
  5. 5.Ecological Monitoring & Research CenterTibetan Environment Monitoring StationLhasaChina

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