Effects of mixed tree species on springtails (Collembola) increase under reduced water availability

Yakun Zhang , Sai Peng , Chen Chen , Han Y. H. Chen , Xinli Chen

Journal of Forestry Research ›› 2026, Vol. 37 ›› Issue (1) : 7

PDF
Journal of Forestry Research ›› 2026, Vol. 37 ›› Issue (1) :7 DOI: 10.1007/s11676-025-01947-3
Original Paper
research-article

Effects of mixed tree species on springtails (Collembola) increase under reduced water availability

Author information +
History +
PDF

Abstract

Soil fauna are crucial for nutrient cycling and promoting plant growth. Plant species mixtures can enhance soil biodiversity and ecosystem functions, but their effects on soil fauna under changing water availability remain poorly understood. To address this gap, we combined a field experiment with a meta-analysis to examine how plant species mixtures influence springtail communities under varying water availability. In a field experiment in Ontario, Canada, we assessed springtail abundance, species richness, Simpson’s diversity index, and community composition in pure and mixed stands of trembling aspen (Populus tremuloides) and jack pine (Pinus banksiana) under ambient, reduced (− 25%), and increased (+ 25%) throughfall in young boreal forest. Tree mixtures enhanced springtail abundance and increased Simpson’s diversity index from − 8.3% under ambient water to + 12.3% under reduced water. Springtail community compositions varied significantly among stand types, with shifts in community composition strongly correlated with fine-root biomass and soil water content. A meta-analysis revealed the effects of plant mixtures on springtail abundance were more positive in sites with less precipitation. On the basis of these results, converting plant mixtures to monocultures will significantly decrease springtail abundance and diversity in areas with less water.

Keywords

Climate change / Water availability / Plant mixture / Springtail / Soil biodiversity

Cite this article

Download citation ▾
Yakun Zhang, Sai Peng, Chen Chen, Han Y. H. Chen, Xinli Chen. Effects of mixed tree species on springtails (Collembola) increase under reduced water availability. Journal of Forestry Research, 2026, 37(1): 7 DOI:10.1007/s11676-025-01947-3

登录浏览全文

4963

注册一个新账户 忘记密码

References

[1]

A’Bear AD, Boddy L, Jones TH. Bottom-up determination of soil Collembola diversity and population dynamics in response to interactive climatic factors. Oecologia, 2013, 173(3): 1083-1087

[2]

Anderson MJ. A new method for non-parametric multivariate analysis of variance. Austral Ecol, 2001, 26(1): 32-46

[3]

Bardgett RD, van der Putten WH. Belowground biodiversity and ecosystem functioning. Nature, 2014, 515(7528): 505-511

[4]

Barrios E. Soil biota, ecosystem services and land productivity. Ecol Econ, 2007, 64(2): 269-285

[5]

Bates D, Maechler M, Bolker B, Walker S (2014) lme4: Linear mixed-effects models using Eigen and S4. R package version 1.1–7

[6]

Blankinship JC, Niklaus PA, Hungate BA. A meta-analysis of responses of soil biota to global change. Oecologia, 2011, 165(3): 553-565

[7]

Bonfanti J, Hedde M, Cortet J, Krogh PH, Larsen KS, Holmstrup M. Communities of collembola show functional resilience in a long-term field experiment simulating climate change. Pedobiologia, 2022, 90 150789

[8]

Bruckner A. Methods for the extraction of microarthropods from soil: a bibliography and guide to the literature. Soil Org, 2024, 96(3): 195-207

[9]

Bruckner A, Barth G, Scheibengraf M. Composite sampling enhances the confidence of soil microarthropod abundance and species richness estimates. Pedobiologia, 2000, 44(1): 63-74

[10]

Canarini A, Fuchslueger L, Schnecker J, Metze D, Nelson DB, Kahmen A, Watzka M, Pötsch EM, Schaumberger A, Bahn M, Richter A. Soil fungi remain active and invest in storage compounds during drought independent of future climate conditions. Nat Commun, 2024, 15(1): 10410

[11]

Chen HYH, Brant AN, Seedre M, Brassard BW, Taylor AR. The contribution of litterfall to net primary production during secondary succession in the boreal forest. Ecosystems, 2017, 20(4830-844

[12]

Chen XL, Chen HYH, Chen C, Peng S. Water availability regulates negative effects of species mixture on soil microbial biomass in boreal forests. Soil Biol Biochem, 2019, 139 107634

[13]

Chen XL, Chen HYH, Searle EB, Chen C, Reich PB. Negative to positive shifts in diversity effects on soil nitrogen over time. Nat Sustain, 2021, 4(3): 225-232

[14]

Chen XL, Peng S, Chen C, Chen HYH. Water availability regulates tree mixture effects on total and heterotrophic soil respiration: a three-year field experiment. Geoderma, 2021, 402 115259

[15]

Crins W, Gray PA, Uhlig P, Wester MC (2009) The ecosystems of Ontario, Part 1: Ecozones and Ecoregions. Inventory, Monitoring and Assessment, SIB TER IMA TR-01

[16]

Escribano-Álvarez P, Pertierra LR, Martínez B, Chown SL, Olalla-Tárraga . Half a century of thermal tolerance studies in springtails (Collembola): a review of metrics, spatial and temporal trends. Curr Res Insect Sci, 2022, 2 100023

[17]

Fick SE, Hijmans RJ. Worldclim 2: new 1-km spatial resolution climate surfaces for global land areas. Int J Climatol, 2017, 37(12): 4302-4315

[18]

Fjellberg A (2010) Corrections and additions to: the Collembola of fennoscandia and Denmark. Part I: poduromorpha.–fauna entomologica Scandinavia, vol 35. In: The Collembola of fennoscandia and Denmark, part II: entomobryomorpha and symphypleona. BRILL, pp 262–263. https://doi.org/10.1163/ej.9789004157705.i-265.28

[19]

Frostegård A, Bååth E. The use of phospholipid fatty acid analysis to estimate bacterial and fungal biomass in soil. Biol Fertil Soils, 1996, 22(1): 59-65

[20]

Ganault P, Nahmani J, Hättenschwiler S, Gillespie LM, David JF, Henneron L, Iorio E, Mazzia C, Muys B, Pasquet A, Prada-Salcedo LD, Wambsganss J, Decaëns T. Relative importance of tree species richness, tree functional type, and microenvironment for soil macrofauna communities in European forests. Oecologia, 2021, 196(2): 455-468

[21]

George PBL, Keith AM, Creer S, Barrett GL, Lebron I, Emmett BA, Robinson DA, Jones DL. Evaluation of mesofauna communities as soil quality indicators in a national-level monitoring programme. Soil Biol Biochem, 2017, 115: 537-546

[22]

Goncharov AA, Leonov VD, Rozanova OL, Semenina EE, Tsurikov SM, Uvarov AV, Zuev AG, Tiunov AV. A meta-analysis suggests climate change shifts structure of regional communities of soil invertebrates. Soil Biol Biochem, 2023, 181 109014

[23]

Gruss I, Twardowski J, Karczewska A, Szopka K, Kluczek K, Magiera-Dulewicz J. Collembola reduce their body sizes under arsenic contamination in the soil–possible use of new screening tool in ecotoxicology. Ecol Indic, 2022, 142 109185

[24]

Hillebrand H, Bennett DM, Cadotte MW. Consequences of dominance: a review of evenness effects on local and regional ecosystem processes. Ecology, 2008, 89(6): 1510-1520

[25]

Holmstrup M. The ins and outs of water dynamics in cold tolerant soil invertebrates. J Therm Biol, 2014, 45: 117-123

[26]

Holmstrup M, Sørensen JG, Schmidt IK, Nielsen PL, Mason S, Tietema A, Smith AR, Bataillon T, Beier C, Ehlers BK. Soil microarthropods are only weakly impacted after 13 years of repeated drought treatment in wet and dry heathland soils. Soil Biol Biochem, 2013, 66: 110-118

[27]

Hopkin SP. Biology of the springtails: (Lnsecta: Collembola), 1997Oxford University Press

[28]

IPCC. Intergovernmental Panel on Climate Change: Climate change 2022—mitigation of climate change: working group III contribution to the sixth assessment report of the intergovernmental panel on climate change, 2023Cambridge University Press

[29]

IPCC. Intergovernmental panel on climate change: climate change 2014: mitigation of climate change. Working group III contribution to the IPCC fifth assessment report., 2015, NY, USA, Cambridge University Press

[30]

Joimel S, Chassain J, Artru M, Faburé J. Collembola are among the most pesticide-sensitive soil fauna groups: a meta-analysis. Environ Toxicol Chem, 2022, 41(10): 2333-2341

[31]

Kaneko N, Salamanca E. Mixed leaf litter effects on decomposition rates and soil microarthropod communities in an oak–pine stand in Japan. Ecol Res, 1999, 14(2): 131-138

[32]

Kardol P, Reynolds WN, Norby RJ, Classen AT. Climate change effects on soil microarthropod abundance and community structure. Appl Soil Ecol, 2011, 47(1): 37-44

[33]

Korboulewsky N, Heiniger C, De Danieli S, Brun JJ. Effect of tree mixture on Collembola diversity and community structure in temperate broadleaf and coniferous forests. For Ecol Manage, 2021, 482 118876

[34]

Korboulewsky N, Perez G, Chauvat M. How tree diversity affects soil fauna diversity: a review. Soil Biol Biochem, 2016, 94: 94-106

[35]

Li ZP, Shi LL, Kuzyakov Y, Pausch J, Scheu S, Pollierer MM. The flux of root-derived carbon via fungi and bacteria into soil microarthropods (Collembola) differs markedly between cropping systems. Soil Biol Biochem, 2021, 160 108336

[36]

Lindberg N, Engtsson JB, Persson T. Effects of experimental irrigation and drought on the composition and diversity of soil fauna in a coniferous stand. J Appl Ecol, 2002, 39(6): 924-936

[37]

Liu HB, Yang LP, Gao JQ, Li QW, Li XL, Feng JG, Yu FH. Increasing plant diversity enhances soil organic carbon storage in typical wetlands of northern China. Front Plant Sci, 2024, 15: 1467621

[38]

Loreau M, Hector A. Partitioning selection and complementarity in biodiversity experiments. Nature, 2001, 412(6842): 72-76

[39]

Ma SJ, Wang QC, Zhang Y, Yan LM, Cui DH, Xu LQ. Effects of natural forest conversion and plantation tree species composition on soil macrofauna communities in Northeast China mountains. J Forestry Res, 2023, 34(51475-1489

[40]

Madej G, Barczyk G, and Gawenda I (2011) Importance of microhabitats for preservation of species diversity, on the basis of mesostigmatid mites (Mesostigmata, Arachnida, Acari). Pol J Environ Stud 20(4)

[41]

Maestre FT, Callaway RM, Valladares F, Lortie CJ. Refining the stress-gradient hypothesis for competition and facilitation in plant communities. J Ecol, 2009, 97(2): 199-205

[42]

Maestre FT, Delgado-Baquerizo M, Jeffries TC, Eldridge DJ, Ochoa V, Gozalo B, Quero JL, García-Gómez M, Gallardo A, Ulrich W, Bowker MA, Arredondo T, Barraza-Zepeda C, Bran D, Florentino A, Gaitán J, Gutiérrez JR, Huber-Sannwald E, Jankju M, Mau RL, Miriti M, Naseri K, Ospina A, Stavi I, Wang DL, Woods NN, Yuan X, Zaady E, Singh BK. Increasing aridity reduces soil microbial diversity and abundance in global drylands. Proc Natl Acad Sci U S A, 2015, 112(51): 15684-15689

[43]

Marx MT, Guhmann P, Decker P. Adaptations and predispositions of different middle European arthropod taxa (Collembola, Araneae, Chilopoda, Diplopoda) to flooding and drought conditions. Animals, 2012, 2(4): 564-590

[44]

Nielsen UN. Climate change impacts on soil fauna, Soil Fauna Assemblages: Global to Local Scales, 2019, Cambridge, Cambridge University Press221245

[45]

Nielsen UN, Osler GHR, Campbell CD, Burslem DFRP, van der Wal R. The influence of vegetation type, soil properties and precipitation on the composition of soil mite and microbial communities at the landscape scale. J Biogeogr, 2010, 37(7): 1317-1328

[46]

Peguero G, Sol D, Arnedo M, Petersen H, Salmon S, Ponge JF, Maspons J, Emmett B, Beier C, Schmidt IK, Tietema A, De Angelis P, Kovács-Láng E, Kröel-Dulay G, Estiarte M, Bartrons M, Holmstrup M, Janssens IA, Peñuelas J. Fast attrition of springtail communities by experimental drought and richness-decomposition relationships across Europe. Glob Chang Biol, 2019, 25(8): 2727-2738

[47]

Peng Y, Peñuelas J, Vesterdal L, Yue K, Peguero G, Fornara DA, Heděnec P, Steffens C, Wu FZ. Responses of soil fauna communities to the individual and combined effects of multiple global change factors. Ecol Lett, 2022, 25(9): 1961-1973

[48]

Podzikowski LY, Heffernan MM, Bever JD. Plant diversity and grasses increase root biomass in a rainfall and grassland diversity manipulation. Front Ecol Evol, 2023, 11: 1259809

[49]

Pretzsch H, del Río M, Schütze G, Ammer C, Annighöfer P, Avdagic A, Barbeito I, Bielak K, Brazaitis G, Coll L, Drössler L, Fabrika M, Forrester DI, Kurylyak V, Löf M, Lombardi F, Matović B, Mohren F, Motta R, den Ouden J, Bravo-Oviedo A. Mixing of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) enhances structural heterogeneity, and the effect increases with water availability. For Ecol Manag, 2016, 373: 149-166

[50]

Qu Q, Wang Z, Gan Q, Liu RT, Xu HW. Impact of drought on soil microbial biomass and extracellular enzyme activity. Front Plant Sci, 2023, 14: 1221288

[51]

Quideau SA, McIntosh ACS, Norris CE, Lloret E, Swallow MJB, Hannam K. Extraction and analysis of microbial phospholipid fatty acids in soils. J vis Exp, 2016, 114 e54360

[52]

R Core Team (2025) R: a language and environment for statistical computing. Version 4.5.0. R Foundation for Statistical Computing

[53]

Salamon JA, Alphei J. The Collembola community of a Central European forest: influence of tree species composition. Eur J Soil Biol, 2009, 45(3): 199-206

[54]

Sánchez-Galindo LM, Sandmann D, Marian F, Krashevska V, Maraun M, Scheu S. Leaf litter identity rather than diversity shapes microbial functions and microarthropod abundance in tropical montane rainforests. Ecol Evol, 2021, 11(52360-2374

[55]

Santonja M, Fernandez C, Proffit M, Gers C, Gauquelin T, Reiter IM, Cramer W, Baldy V. Plant litter mixture partly mitigates the negative effects of extended drought on soil biota and litter decomposition in a Mediterranean oak forest. J Ecol, 2017, 105(3): 801-815

[56]

Soong JL, Vandegehuchte ML, Horton AJ, Nielsen UN, Denef K, Shaw EA, de Tomasel CM, Parton W, Wall DH, Cotrufo MF. Soil microarthropods support ecosystem productivity and soil C accrual: evidence from a litter decomposition study in the tallgrass prairie. Soil Biol Biochem, 2016, 92: 230-238

[57]

Tilman D. Resource competition and community structure. Monogr Popul Biol, 1982, 17: 1-296

[58]

Turnbull MS, Lindo Z. Combined effects of abiotic factors on Collembola communities reveal precipitation may act as a disturbance. Soil Biol Biochem, 2015, 82: 36-43

[59]

Turnbull MS, Stebaeva S. Collembola of Canada Zookeys, 2019, 819: 187-195

[60]

Ullah S, Wu JP, Ali Shah J, Wang XM, Lyu YM, Guo ZW, Ali K, Chen DY, Sun H. Tree diversity drives understory carbon storage rather than overstory carbon storage across forest types. J Forestry Res, 2024, 35(1): 125

[61]

Wan XH, Joly FX, Jia H, Zhu M, Fu YR, Huang ZQ. Functional identity drives tree species richness-induced increases in litterfall production and forest floor mass in young tree communities. New Phytol, 2023, 240(31003-1014

[62]

Wang Y, Slotsbo S, Holmstrup M. Soil dwelling springtails are resilient to extreme drought in soil, but their reproduction is highly sensitive to small decreases in soil water potential. Geoderma, 2022, 421 115913

[63]

Wise DH, Lensing JR. Impacts of rainfall extremes predicted by climate-change models on major trophic groups in the leaf litter arthropod community. J Anim Ecol, 2019, 88(10): 1486-1497

[64]

Wright AJ, Ebeling A, de Kroon H, Roscher C, Weigelt A, Buchmann N, Buchmann T, Fischer C, Hacker N, Hildebrandt A, Leimer S, Mommer L, Oelmann Y, Scheu S, Steinauer K, Strecker T, Weisser W, Wilcke W, Eisenhauer N. Flooding disturbances increase resource availability and productivity but reduce stability in diverse plant communities. Nat Commun, 2015, 6: 6092

[65]

Wright AJ, Mommer L, Barry K, van Ruijven J. Stress gradients and biodiversity: monoculture vulnerability drives stronger biodiversity effects during drought years. Ecology, 2021, 102(1 e03193

[66]

Xi NX, Chen DX, Liu W, Bloor JMG. Positive plant diversity effects on soil microbial drought resistance are linked to variation in labile carbon and microbial community structure. Funct Ecol, 2023, 37(9): 2347-2357

[67]

Xu GL, Kuster TM, Günthardt-Goerg MS, Dobbertin M, Li MH. Seasonal exposure to drought and air warming affects soil collembola and mites. PLoS ONE, 2012, 7(8 e43102

[68]

Yachi S, Loreau M. Biodiversity and ecosystem productivity in a fluctuating environment: the insurance hypothesis. Proc Natl Acad Sci USA, 1999, 96(41463-1468

[69]

Zagatto MRG, de Araujo Pereira AP, de Souza AJ, Pereira RF, Baldesin LF, Pereira CM, Lopes RV, Cardoso EJBN. Interactions between mesofauna, microbiological and chemical soil attributes in pure and intercropped Eucalyptus grandis and Acacia mangium plantations. For Ecol Manage, 2019, 433: 240-247

[70]

Zhang HY, Han GX, Huang TS, Feng Y, Tian W, Wu XC. Mixed forest of Larix principis-rupprechtii-rupprechtii and Betula platyphylla modulating soil fauna diversity and improving faunal effect on litter decomposition. Forests, 2022, 13(5): 703

[71]

Zhang Y, Chen HYH, Taylor AR. Positive species diversity and above-ground biomass relationships are ubiquitous across forest strata despite interference from overstorey trees. Funct Ecol, 2017, 31(2419-426

[72]

Zhang YK, Peng S, Chen XL, Chen HYH. Plant diversity increases the abundance and diversity of soil fauna: a meta-analysis. Geoderma, 2022, 411 115694

[73]

Zhang YK, Peng S, Ma ZL, Chen C, Gao BL, Chen XL, Chen HYH. Increased positive tree species mixture effects on the abundance and richness of Collembola with stand development in Canadian boreal forests. For Ecosyst, 2025, 13 100294

[74]

Zheng LT, Chen HYH, Yan ER. Tree species diversity promotes litterfall productivity through crown complementarity in subtropical forests. J Ecol, 2019, 107(4): 1852-1861

[75]

Zhou ZH, Wang CK, Luo YQ. Meta-analysis of the impacts of global change factors on soil microbial diversity and functionality. Nat Commun, 2020, 11(13072

[76]

Zhu XW, Fu ZG, Gong HD, Kuzyakov Y, Lu ZY, Zhang J, Zhu XM, Hong SS, Song QH, Zhang YP, Wen HD, Zhou WJ. Drought effects on litter fraction and recovery in a subtropical forest. Ecol Indic, 2024, 166 112429

RIGHTS & PERMISSIONS

The Author(s)

PDF

30

Accesses

0

Citation

Detail

Sections
Recommended

/