Leaf litter decomposition in Pinus massoniana monoculture and mixed plantation with Castanopsis fissa in a subtropical area of China

Na Lin; Huangqiang Huang; Zhanpeng Qiu; Zhifei Li; Yuheng He; Tao Ma; Shiqing Chen

doi:10.15287/afr.2026.3674

Authors

Na Lin ‌College of Forestry and Landscape Architecture, South China Agricultural University, Guangdong, China
Huangqiang Huang ‌College of Forestry and Landscape Architecture, South China Agricultural University, Guangdong, China
Zhanpeng Qiu ‌College of Forestry and Landscape Architecture, South China Agricultural University, Guangdong, China
Zhifei Li ‌College of Forestry and Landscape Architecture, South China Agricultural University, Guangdong, China
Yuheng He ‌College of Forestry and Landscape Architecture, South China Agricultural University, Guangdong, China
Tao Ma ‌College of Forestry and Landscape Architecture, South China Agricultural University, Guangdong, China
Shiqing Chen ‌College of Forestry and Landscape Architecture, South China Agricultural University, Guangdong, China

DOI:

https://doi.org/10.15287/afr.2026.3674

Keywords:

mixed leaf litter, decomposition, coniferous monoculture, broadleaved species, ubtropical area

Abstract

We measured the rates of Pinus massoniana needle litter decomposition mixed with different proportions of Castanopsis fissa leaf litter in P. massoniana monoculture and P. massoniana mixed with C. fissa forests, which are the typical forest of the subtropical zone of China. The broadleaved species on sites could naturally promote the degradation of litter and mineralization of carbon and nitrogen, which results in faster nutrient return and enhanced productivity. But it is not well known how specific species influence the decomposition processes of each other in mixtures. In this paper, there were 4 types of litter mixtures, which referred to the mass ratio of P. massoniana leaf to C. fissa leaf in the mixtures: 1:1, 3:2, 7:3, and 1:0. We placed 4 types of litterbags in the two forest types for an over 12-month experiment. We applied the litterbag technique for this experiment. The main results were: 1. P. massoniana leaf litter (1:0) decomposed faster in the mixed forest than in monoculture. 2. The mixed leaf litter (1:1, 3:2, 7:3) decomposed faster than the monoculture P. massoniana litter, moreover, the 1:1 type decomposed the fastest. 3. The mixture effects on needle litter decomposition changed during decomposition process, the antagonistic mixture effects of 1:1 and 7:3 could be found in the first six months in the mixed and monoculture stands, the non-additive synergistic litter mixture effects dominated after 9 months. The results showed that broadleaff litter could promote the decomposition of conifer litter, and the decomposition of litter was faster in the mixed forest than in the monoculture coniferous forest. It also shows that keeping a suitable proportion of broad-leaved tree species in coniferous monoculture forest is beneficial to the nutrient cycle and the tree’s growth.

References

Aber J.D., Melillo J.M., Nadelhoffer K.J., Pastor J., Boone R.D., 1991. Factors controlling nitrogen cycling and nitrogen saturation in Northern temperate forest ecosystems. Ecological Applications 1(3): 303-315. https://doi.org/10.2307/1941759

Aber J.D., Mellilo J.M., 1991. Terrestrial ecosystems. Saunders College Publishing, Philadelphia, PA.

Ágoston-Szabó E., Schöll K., Kiss A., Dinka M., 2017. The effects of tree species richness and composition on leaf litter decomposition in a Danube oxbow lake (Gemenc, Hungary). Fundamental & Applied Limnology 189(4): 301-314. https://doi.org/ 10.1127/fal/2017/0675

Ayres E., Steltzer H., Simmons B.L., Simpson R.T., Steinweg J.M., Wallenstein M.D., Mellor N., Parton W.J., Moore J.C., Wall D.H., 2009. Home-field advantage accelerates leaf litter decomposition in forests. Soil Biology & Biochemistry 41: 606–610. https://doi.org/10.1016/j.soilbio.2008.12.022

Bai Y.X., Zhou Y.C., Chen X.L., An Z.F., Zhang X.Y., Du J.J., Chang S.X., 2023. Tree species composition alters the decomposition of mixed litter and the associated microbial community composition and function in subtropical plantations in China. Forest Ecology and Management 529: 120743. https://doi.org/10.1016/j.foreco.2022.120743

Berg B., 2000. Litter decomposition and organic matter turnover in northern f10.1016/S0378-1127(99)00294-7orest soils. Forest Ecology and Management 133: 13-22. https://doi.org/ 10.1016/S0378-1127(99)00294-7

Berg B., Ekbohm G., Söderström B., Staaf H., 1991. Reduction of decomposition rates of Scots pine needle litter due to heavy-metal pollution. Water Air & Soil Pollution 59(1-2): 165-177. https://doi.org/10.1007/BF00283179

Berg B., McClaugherty, C., 2008. Plant Litter. Springer-Verlag, Berlin. https://doi.org/10.1007/978-3-540-74923-3_7

Berg B., Staaf H., 1980. Decomposition rate and chemical changes of Scots pine needle litter. II. Influence of chemical composition. In: T Persson (ed) Structure and function of northern coniferous forests. An ecosystem study. Ecological Bulletins, Stockholm 33, pp 163–178.

Bhatnagar J.M., Peay K.G., Treseder K.K., 2018. Litter chemistry influences decomposition through activity of specific microbial functional guilds. Ecological Monographs 88: 429–444. https://doi.org/10.1002/ecm.1303.

Bocock K.L., Gilbert J.W., 1957. The disappearance of leaf litter under different woodland conditions. Plant & Soil 9 (2): 179–185. https://doi.org/10.1007/BF01398924.

Bonanomi G., Incerti G., Antignani V., Capodilupo M., Mazzoleni S., 2010. Decomposition and nutrient dynamics in mixed litter of Mediterranean species. Plant and Soil 331: 481-496. https://doi.org/10.1007/s11104-009-0269-6

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 Biology & Biochemistry 77: 121-128. https://doi.org/10.1016/j.soilbio.2014.06.019.

Cardinale B.J., Matulich K.L., Hooper D.U., Byrnes J.E., Duffy E., Gamfeldt L., Balvanera P., O'Connor M.I., Gonzalez A., 2011. The functional role of producer diversity in ecosystems. American Journal of Botany 98(3): 572–592. https://doi.org/10.3732/ajb.1000364.

Chapin I.F.S., Matson P.A., Vitousek P., 2011. Principles of terrestrial ecosystem ecology, New York, NY: Springer.

Chapman S.K., Koch G.W., 2007. What type of diversity yields synergy during mixed litter decomposition in a natural forest ecosystem? Plant and Soil 299: 153-162. https://doi.org/10.1007/s11104-007-9372-8

Chapman S.K., Newman G.S., 2010. Biodiversity at the plant-soil interface: microbial abundance and community structure respond to litter mixing. Oecalogia 162: 763-769. https://doi.org/10.1007/s00442-009-1498-3

Chen B.M., Peng S.L., D'Antonio C.M., Li D.J., Ren W.T., 2013. Non-additive effects on decomposition from mixing litter of the invasive Mikania micrantha HBK with native plants. PloS ONE 8(6): e66289. https://doi.org/10.1371/journal.pone.0066289.

Chomel M., Guittonny-Larchevȇque M., Fernandez C., Gallet C., DesRochers A., Parė D., Jackson B.G., Baldy V., 2016. Plant secondary metabolites: a key driver of litter decomposition and soil nutrient cycling. Journal of Ecology 104(6): 1-15. https://doi.org/10.1111/1365-2745.12644

Cuchietti A., Marcotti E., Gurvich D.E., Cingolani A.M., Harguindeguy N.P., 2014. Leaf litter mixtures and neighbour effects: Low-nitrogen and high-lignin species increase decomposition rate of high-nitrogen and low-lignin neighbours. Applied Soil Ecology 82: 44-51. https://doi.org/10.1016/j.apsoil.2014.05.004

Djukic I., Kepfer-Rojas S., Schmidt I.K., Larsen K.S., Beier C., Berg B., Verheyen K., 2018. Early stage litter decomposition across biomes. Science of the Total Environment 628–629: 1369–1394. https://doi.org/10.1016/j.scitotenv.2018.01.012.

Dong M., Wang Y.F., Kong F.Z., Jiang G.M., Zhang Z.B., 1996. Observation and Analysis of Terrestrial Biocommunities. Standard Press of China, Beijing.

Fang H., Mo J., Peng S., Li Z., Wang H., 2007. Cumulative effects of nitrogen additions on litter decomposition in three tropical forests in southern China. Plant and Soil 297:233–242. https://doi.org/10.1007/s11104-007-9339-9

García-Palacios P., Shaw E.A., Wall D.H., Hättenschwiler S., Laliberté E., 2017. Contrasting mass-ratio vs. niche complementarity effects on litter C and N loss during decomposition along a regional climatic gradient. Journal of Ecology 105 (4): 968–978. https://doi.org/10.1111/1365-2745.12730

Gartner T.B., Cardon Z.G., 2004. Decomposition dynamics in mixed-species leaf litter. Oikos 104: 230-246. https://doi.org/10.1111/j.0030-1299.2004.12738.x.

Gholz H.L., Wedin D.A., Smitherman S.M., Harmon M.E., Parton W.J., 2000. Longterm dynamics of pine and hardwood litter in contrasting environments: toward a global model of decomposition. Global Change Biology 6: 751–765. https://doi.org/10.1046/j.1365-2486.2000.00349.x

Handa I.T., Aerts R., Berendse F., Berg M.P., Bruder A., Butenschoen O., Chauvet E., Gessner M.O., Jabiol J., Makkonen M., 2014. Consequences of biodiversity loss for litter decomposition across biomes. Nature 509(7499): 218–221. https://doi.org/10.1038/nature13247.

Hättenschwiler S., Gasser P., 2005. Soil animals alter plant litter diversity effects on decomposition. Proceedings of the National Academy of Sciences of the United States of America 102(5): 1519-1524. https ://doi.org/10.1073/pnas.0404977102.

He Z.M., Yu Z.P., Huang Z.Q., Davis M., Yang Y.S., 2016. Litter decomposition, residue chemistry and microbial community structure under two subtropical forest plantations: A reciprocal litter transplant study. Applied Soil Ecology 101: 84-92. https://doi.org/10.1016/j.apsoil.2016.01.015.

Hector A., Beale A.J., Minns A., Otway S.J., Lawton J.H., 2000. Consequences of the reduction of plant diversity for litter decomposition: effects through litter quality and microenvironment. OIKOS 90: 357-371. https://doi.org/10.1034/j.1600-0706.2000.900217.x.

Heinrichs H., Brumsack H.J., Loftfield N., König N., 1986. Verbessertes Druckaufschlußsz Tellerradgehaeutem für biologische und anorganische Materialien. Journal of Plant Nutriention and Soil Science 149: 350-353.

Hobbie S.E., 2006. Contrasting effects of substrate and fertilizer nitrogen on the early stages of litter decomposition. Ecosystems 8: 644-656. https://doi.org/10.1007/s10021-003-0110-7

Hooper D.U., Chapin F.S., Ewel J.J., Hector A., Inchausti P., Lavorel S., Lawton J.H., Lodge D.M., Loreau M., Naeem S., Schmid B., Setala H., Symstad A.J., Vandermeer L., Wardle D.A., 2005. Effects of biodiversity on ecosystem functioning: a consensus of current knowledge. Ecological Monographs 75(1): 3–35. https://doi.org/10.1890/04-0922.

Kaspari M., Garcia M.N., Harms K.E., Santana M., Wright S.J., Yavitt J.B., 2008. Multiple nutrients limit litter fall and decomposition in a tropical forest. Ecology Letters 11: 35–43. https://doi.org/10.1111/j.1461-0248.2007.01124.

Lecerf A., Marie G., Kominoski J.S., LeRoy C.J., Bernadet C., Swan C.M., 2011. Incubation time, functional litter diversity, and habitat characteristics predict litter-mixing effects on decomposition. Ecology 92: 160-169. https://doi.org/10.2307/29779584.

Li S.S., Tong Y.W., Wang Z.W., 2017. Species and genetic diversity affect leaf litter decomposition in subtropical broadleaved forest in southern China. Journal of Plant Ecology 10(1): 232-241. https://doi.org/10.1093/jpe/rtw122

Li Y.H., Zhou L., Wu J., Zhou W.M., Dai L.M., Lu Z.M., Huang L.Y., 2017. Decomposition of mixed leaf litter of Larix kaempferi plantation in eastern montane region of Liaoning Province. Chinese Journal of Ecology 36(11): 3049-3055. (in Chinese)https ://doi.org/10.13292/j.1000-4890.201711.037

Lin N., Barsch N., Heinrichs S., Vor, T., 2015. Long-term effects of canopy opening and liming on leaf litter production, and on leaf litter and fine-root decomposition in a European beech (Fagus sylvatica) forest. Forest Ecology and Management 338: 183–190. https://doi.org/10.1016/j.foreco.2014.11.029.

Lin N., Xiao L.L., Qiu Z.P., Li Z.F., He J.L., Ma T., Chen S.Q., 2025. Spatiotemporal variation and driving mechanisms of planted forest biomass carbon in Guangdong south China. Environmental and Sustainability Indicatior 28: 100990. https://doi.org/10.1016/j.indic.2025.100990

Liu P., Sun O.J., Huang J.H., Li L.H., Han X.G., 2007. Nonadditive effects of litter mixtures on decomposition and correlation with initial litter N and P concentrations in grassland plant species of northern China. Biology and Fertility of Soil 44 (1): 211-216. https://doi.org/10.1007/s00374-007-0195-9

Liu S.Q., Yang R., Peng X.D., Hou C.L., Ma J.B., Guo J.R., 2022. Contributions of plant litter decomposition to soil nutrients in ecological tea gardens. Agriculture 12(7): 957. https://doi.org/10.3390/agriculture12070957

Makkonen M., Berg M.P.B., van Logtestijn R.S.P., van Hal J.R., Aerts R., 2012. Do physical plant litter traits explain non-additivity in litter mixtures? A test of the improved microenvironmental conditions theory. Oikos 122: 987-997. https://doi.org/10.1111/j.1600-0706.2012.20750.x

Meiwes K.J.; Khanna P.K., Ulrich B., 1986. Parameters for describing soil acidification and their relevance to stability of forest ecosystems. Forest Ecology and management 15: 161–179. https://doi.org/10.1016/0378-1127(86)90064-2

Melillo J.M., Aber J.D., Muratore J.F., 1982. Nitrogen and lignin control of hardwood leaf litter decomposition dynamics. Ecology 63(3): 621-626. https://doi.org/10.1007/s10021-017-0130-3.

Mládková P., Mládek J., Hejduk S., Hejcman M., Pakeman R., 2018. Calcium plus magnesium indicates digestibility: the significance of the second major axis of plant chemical variation for ecological processes. Ecology Letters 21 (6): 885–895. https://doi.org/10.1111/ele.12956.

Mo J.M., Brown S., Kong G.H., Lenart M., Zhang Y.C., 1996. Litter decomposition and its nutrient dynamics of a Pinus massoniana forest in Dinghushan Biosphere Reserve. Acta Phytoecologica Sinica 20(6): 534-542.

Montané F., Romanyà J., Rovira P., Casals P., 2013. Mixtures with grass litter may hasten shrub litter decomposition after shrub encroachment into mountain grasslands. Plant and Soil 368(1-2): 459-469. https://doi.org/10.1007/s11104-012-1533-8

Olson J.S., 1963. Energy storage and the balance of producers and decomposers in ecological systems. Ecology 44(2): 322–331.

Pan J.W., Li J.Y., He Q., Su Y., Liu X.D., Qiu Q., Lin N., 2020. Study on the soil nutrient characteristic and their relationship with litter quality in two forest types of Pinus massoniana. Journal of Central South University of Forestry & Technology 40(2): 54-63. (in Chineses).

Porre R.J., van der Werf W., De Deyn G.B., Stomph T.J., 2020. Is litter decomposition enhanced in species mixtures? A meta-analysis. Soil Biology & Biochemistry 145: 107791. https://doi.org/10.1016/j.soilbio.2020.107791

Prentice I.C., Farquhar G., Fasham M., Goulden M., Heimann M., Jaramillo V.J., Kheshgi H., Quéré C.L., Scholes R., Wallace, D., 2001. The carbon cycle and atmospheric carbon dioxide. Cambridge, UK: Cambridge University Press.

Prescott C.E., 2024. Perspectives: Regenerative forestry - Managing forests for soil life. Forest Ecology and Management 554: 121674. https://doi.org/10.1016/j.foreco.2023.121674

Prescott C.E., Blevins L.L., Staley C., 2004. Litter decomposition in British Columbia forests: controlling factors and influences of forestry activities. BC Journal of Ecosystems and Management 5: 44–57. https://doi.org/10.22230/jem.2005v5n2a298.

Prescott C.E., Reid A., Wu S.Y., Nilsson M.C., 2017. Decomposition rates of surface and buried forest-floor material. Canadian Journal of Forest Research 47(8): 1140-1144. https://doi.org/10.1139/cjfr-2016-0533

Prescott C.E., Zabek L.M., Staley C.L., Kabzems R., 2000. Decomposition of broadleaf and needle litter in forests of British Columbia: influences of litter type, forest type, and litter mixtures. Canadian Journal of Forest Research 30: 1742-1750. https://doi.org/10.1139/cjfr-30-11-1742

Schaefer D., Steinberger Y., Whitford W.G., 1985. The failure of nitrogen and lignin control of decomposition in an North American desert. Oecologia 65: 382–386. https://doi.org/10.1007/BF00378913

Schimel J.P., Hättenschwiler S., 2007. Nitrogen transfer between decomposing leaves of different N status. Soil Biology & Biochemistry 39: 1428-1436. https://doi.org/10.1016/j.soilbio.2006.12.037

Schindler M., Gessner M., 2009. Functional leaf traits and biodiversity effects on litter decomposition in a stream. Ecology 90(6): 1641–1649. https://doi.org/10.1890/08-1597.1

Schlesinger W.H., 1997. Biogeochemistry: an analysis of global change, 2nd ed, Academic Press.

Seatedt T.R., 1984. The role of microarthropods in the decomposition and mineralization of N. Chemical Engineering Science 48(20): 3529-3534. https://doi.org/10.1146/annurev.en.29.010184.000325

Setiawan N.N., Vanhellemont M., Baeten L., Gobin R., De Smedt P., Proesmans W., Ampoorter E., Verheyen K., 2016. Dose neighbourhood tree diversity affect the crown arthropod community in saplings? Biodiversity and Conservation 25(1): 169-185. https://doi.org/10.1007/s10531-015-1044-z

Smith J.S., Valentine K., Small J.M., Khan A., Gedan K., Nordio G., Fagherazzi S., Kirwan M.L., 2024. Litter decomposition in retreating coastal forests. Estuaries and Coasts 47: 1139-1149. https://doi.org/10.1007/s12237-024-01358-3

Srivastava D.S., Cardinale B.J., Downing A., Duffy J.E., Jouseau C., Sankaran M., Wright J.P., 2009. Diversity has stronger top-down than bottom-up effects on decomposition. Ecology, 90(4), 1073-1083. https://doi.org/10.1890/08-0439.1

Staaf H., 1980. Influence of chemical composition, addition of raspberry leaves, and nitrogen supply on decomposition rate and dynamics of nitrogen and phosphorus in beech leaf litter. Oikos 35: 55–62. https://doi.org/10.2307/3544726

Trogisch S., He J.S., 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 and Soil 400: 337-350. https://doi.org/10.1007/s11104-015-2737-5

Vos V.C.A., van Ruijven J., Berg M.P., Peeters E.T.H.M., Berendse F., 2013. Leaf litter quality drives litter mixing effects through complementary resource use among detritivores. Oecologia 173(1): 269-280. https://doi.org/10.1007/s00442-012-2588-1

Wardle D.A., Bonner K.I., Barker G.M., 2002. Linkages between plant litter decomposition, litter quality, and vegetation responses to herbivores. Functional Ecology16: 585-595. https://doi.org/10.1046/j.1365-2435.2002.00659.x

Wardle D.A., Bonner K.I., Nicholson K.S., 1997. Biodiversity and plant litter: experimental evidence which does not support the view that enhanced species richness improves ecosystem function. Oikos 79: 247-258. https://doi.org/10.2307/3546010

Wardle D.A., Nilsson M-C., Zackrisson O., Gallet C., 2003. Determinants of litter mixing effects in a Swedish boreal forest. Soil Biology & Biochemistry 35: 827-835. https://doi.org/10.1016/S0038-0717(03)00118-4.

Wider R.K., Lang G.E., 1982. A critique of the analytical methods used in examining decomposition data obtained from litter bags. Ecology 63: 1636–1642. https://doi.org/10.2307/1940104

Wu D.D., Li T.T., Wan S.Q., 2013. Time and litter species composition affect litter-mixing effects on decomposition rates. Plant and Soil 371: 355-366. https://doi.org/10.1007/s11104-013-1697-x

Yang H.X., Wang S.L., Fan B., Zhang W.D., 2010. Dynamics of annual litter mass and nutrient return of different ages Masson pine plantations. Chinese Journal of Ecology 29(12): 2334-2340.

Zeng X.Y., Gao H.L., Wang R.X., Majcher B.M., Woon J.S., Wenda C., Eggleton P., Griffiths H.M., Ashton L.A., 2024. Global contribution of invertebrates to forest letter decomposition. Ecology Letters 27: e14423. https://doi.org/10.1111/ele.14423

Zhao J.D., Yang Y.C., Shen F.Y., Jun M., Yang L.X., 2024. Dynamic characteristics of litter decomposition and nutrient release in 4 species of Larix plantations. Journal of Central South University of Forestry & Technology 44(7): 119-129. (in Chinese).

Zheng J.Q., Guo R.H., Li D.S., Zhang J.H., Han S.J., 2017. Nitrogen addition, drought and mixture effects on litter decomposition and nitrogen immobilization in a temperate forest. Plant and Soil 416: 165-179. https://doi.org/10.1007/s11104-017-3202-4

Zhou S.X., Butenschoen O., Barantal S., Handa I.T., Makkonen M., Vos V., Aerts R., Berg M.P., McKie B., Ruijven J.V., Hättenschwiler S., Scheu S., 2020. Decomposition of leaf litter mixtures across biomes: The role of litter identity, diversity and soil fauna. Journal of Ecology 108: 2283-2297. https://doi.org/10.5061/dryad.9cnp5hqdk