Long-term spruce canopy gaps reduce soil nematode functional diversity via taxa loss and trait convergence

Chengwei Tu , Wenchao Yan , Yan Zhang , Ajuan Zhang , Yilin Feng , Xueyong Pang

Journal of Forestry Research ›› 2026, Vol. 37 ›› Issue (1) : 133

PDF
Journal of Forestry Research ›› 2026, Vol. 37 ›› Issue (1) :133 DOI: 10.1007/s11676-026-02076-1
Original Paper
research-article
Long-term spruce canopy gaps reduce soil nematode functional diversity via taxa loss and trait convergence
Author information +
History +
PDF

Abstract

Forest gaps are widely implemented to enhance aboveground biodiversity within continuous cover forestry, yet their long-term impacts on soil fauna functional dynamics remain unclear. We used a trait-based framework to reveal how old gaps in subalpine spruce plantations affect the nematode functional identity (integrating body weight, trophic level, and lifespan) and functional richness (multidimensional functional space volume). We found that gaps reduced soil moisture, driving nematode communities toward fast-growing strategies characterized by smaller body weight (weighted mean biomass reduced 58%) and shorter lifespan (proportion of long-lived taxa decreased 66%). Moreover, nematode functional richness decreased by 38% in gaps through dual pathways: reduced taxa richness (relative importance 0.51) linked to litter stoichiometry; and moisture-regulated trait convergence (relative importance 0.32, eliminated resource-conserving strategists). Neither nematode functional identity nor functional richness affected soil multifunctionality, highlighting canopy loss may disrupt the nematode-soil functioning relationships. These findings indicate that understory development alone is insufficient to enhance belowground functionality, underscoring the need to align aboveground and belowground restoration goals in forestry practices.

Graphical abstract

Keywords

Forest gap / Planted forest / Soil nematode / Functional trait / Soil multifunctionality

Cite this article

Download citation ▾
Chengwei Tu, Wenchao Yan, Yan Zhang, Ajuan Zhang, Yilin Feng, Xueyong Pang. Long-term spruce canopy gaps reduce soil nematode functional diversity via taxa loss and trait convergence. Journal of Forestry Research, 2026, 37 (1) : 133 DOI:10.1007/s11676-026-02076-1

登录浏览全文

4963

注册一个新账户 忘记密码

References

[1]

Amyntas A, Berti E, Gauzens B, Albert G, Yu WT, Werner AS, Eisenhauer N, Brose U. Niche complementarity among plants and animals can alter the biodiversity–ecosystem functioning relationship. Funct Ecol, 2023, 37(10): 2652-2665.

[2]

Andriuzzi WS, Franco ALC, Ankrom KE, Cui SY, de Tomasel CM, Guan PT, Gherardi LA, Sala OE, Wall DH. Body size structure of soil fauna along geographic and temporal gradients of precipitation in grasslands. Soil Biol Biochem, 2020, 140. ArticleID: 107638

[3]

Arunachalam A, Arunachalam K. Influence of gap size and soil properties on microbial biomass in a subtropical humid forest of north-east India. Plant Soil, 2000, 223(1): 187-195.

[4]

Bardgett RD, van der Putten WH (2014) Belowground biodiversity and ecosystem functioning. Nature 515(7528):505–511. https://doi.org/10.1038/nature13855

[5]

Bonfanti J, Potapov AM, Angst G, Ganault P, Briones MJI, Calderón-Sanou I, Chen TW, Conti E, Degrune F, Eisenhauer N, Ferlian O, Hackenberger D, Hauer A, Hedde M, Hohberg K, Krogh PH, Mulder C, Perez-Roig C, Russell D, Shelef O, Zhou Z, Zuev AG, Berg MP. Linking effect traits of soil fauna to processes of organic matter transformation. Funct Ecol, 2025, 39(2): 446-461.

[6]

Byrnes JEK, Gamfeldt L, Isbell F, Lefcheck JS, Griffin JN, Hector A, Cardinale BJ, Hooper DU, Dee LE, Duffy EJ. Investigating the relationship between biodiversity and ecosystem multifunctionality: challenges and solutions. Meth Ecol Evol, 2014, 5(2): 111-124.

[7]

Clinton BD. Light, temperature, and soil moisture responses to elevation, evergreen understory, and small canopy gaps in the southern Appalachians. For Ecol Manag, 2003, 186(1–3): 243-255.

[8]

DeForest JL. The influence of time, storage temperature, and substrate age on potential soil enzyme activity in acidic forest soils using MUB-linked substrates and L-DOPA. Soil Biol Biochem, 2009, 41(6): 1180-1186.

[9]

Duncker PS, Barreiro SM, Hengeveld GM, Lind T, Mason WL, Ambrozy S, Spiecker H. Classification of forest management approaches: a new conceptual framework and its applicability to European forestry. Ecol Soc, 2012, 17(4. ArticleID: art51

[10]

Eisenhauer N, Hines J, Maestre FT, Rillig MC. Reconsidering functional redundancy in biodiversity research. Npj Biodivers, 2023, 2. ArticleID: 9

[11]

Erktan A, Or D, Scheu S. The physical structure of soil: determinant and consequence of trophic interactions. Soil Biol Biochem, 2020, 148. ArticleID: 107876

[12]

Gao K, Du H, Zhu ZH, Fang YT, Li DJ. Increasing plant species diversity aggravates microbial phosphorus limitation but alleviates microbial carbon limitation in a subtropical forest. J Plant Ecol, 2024, 17(6. ArticleID: rtae100

[13]

Geisen S, Snoek LB, ten Hooven FC, Duyts H, Kostenko O, Bloem J, Martens H, Quist CW, Helder JA, van der Putten WH. Integrating quantitative morphological and qualitative molecular methods to analyse soil nematode community responses to plant range expansion. Methods Ecol Evol, 2018, 9(6): 1366-1378.

[14]

Geisen S, Wall DH, van der Putten WH. Challenges and opportunities for soil biodiversity in the anthropocene. Curr Biol, 2019, 29(19): R1036-R1044.

[15]

Graf M, Achury R, Lanzrein I, Wenglein R, Annighöfer P, Scheu S, Weisser WW. The effect of Douglas-fir on biodiversity in European forests–What do we know and what do we not know?. For Ecosyst, 2025, 13. ArticleID: 100319

[16]

Griffiths HM, Ashton LA, Parr CL, Eggleton P. The impact of invertebrate decomposers on plants and soil. New Phytol, 2021, 231(6): 2142-2149.

[17]

Grime JP. Benefits of plant diversity to ecosystems: immediate, filter and founder effects. J Ecol, 1998, 86(6): 902-910.

[18]

Hayden HL, Ghaderi R, Trollip C, Hu HW, He JZ. Advancing the use of metabarcoding derived nematode-based indices as soil health bioindicators in agricultural and natural environments. Soil Biol Biochem, 2025, 205. ArticleID: 109772

[19]

Hou MT, Zhang GQ, Liu QF, Zhao YQ, Zang LP, Chen DM, Sui MZ, He YJ, Ding FJ. Canopy gaps amplify stochastic assembly and reduce network complexity of soil microbial community in a karst forest. CATENA, 2025, 254. ArticleID: 109009

[20]

Huang YM, Yang X, Zhang DJ, Zhang J. The effects of gap size and litter species on colonization of soil fauna during litter decomposition in Pinus massoniana plantations. Appl Soil Ecol, 2020, 155. ArticleID: 103611

[21]

Inkotte J, Bomfim B, da Silva SC, Valadão MBX, da Rosa MG, Viana RB, Gatto A, Pereira RS. Litter removal impacts on soil biodiversity and eucalypt plantation development in the seasonal tropics. J for Res, 2023, 34(3): 735-748.

[22]

Isaacson BN, Yang Y, Anderson MC, Clark KL, Grabosky JC. The effects of forest composition and management on evapotranspiration in the New Jersey Pinelands. Agric for Meteorol, 2023, 339. ArticleID: 109588

[23]

Jian ZJ, Zeng LX, Lei L, Liu CF, Shen YF, Zhang JJ, Xiao WF, Li MH. Effects of thinning and understory removal on soil phosphorus fractions in subtropical pine plantations. Front Plant Sci, 2024, 15. ArticleID: 1416852

[24]

Jiang Y, Wang ZH, Liu Y, Han YL, Wang Y, Wang Q, Liu T. Nematodes and their bacterial prey improve phosphorus acquisition by wheat. New Phytol, 2023, 237(3): 974-986.

[25]

Jiang YR, Wang Z, Cao R, Wang Q, Li WZ, Yang WQ. Decaying logs and gap positions jointly maintain the structure and function of soil invertebrate community in a subalpine forest. Plant Soil, 2025, 512(1): 977-990.

[26]

Liao XH, Fu SL, Zhao J. Altered energy dynamics of multitrophic groups modify the patterns of soil CO2 emissions in planted forest. Soil Biol Biochem, 2023, 178. ArticleID: 108953

[27]

Liao XH, Zhao J, Magura T, Zhang W, Hu PL, Xiao D, Li JN, Zhang J, Wang KL. Strengthened trophic interactions in soil micro-food webs stimulate soil carbon storage under vegetation restoration. J Appl Ecol, 2025, 62(10): 2594-2611.

[28]

Liu SR, Yang YJ, Wang H (2018) Development strategy and management countermeasures of planted forests in China: transforming from timber-centered single objective management towards multi-purpose management for enhancing quality and benefits of ecosystem services. Acta Ecol Sin 38(1):1–10. https://doi.org/10.5846/stxb201712072201

[29]

Loreau M, Naeem S, Inchausti P, Bengtsson J, Grime JP, Hector A, Hooper DU, Huston MA, Raffaelli D, Schmid B, Tilman D, Wardle DA (2001) Biodiversity and ecosystem functioning: current knowledge and future challenges. Science 294(5543):804–808. https://doi.org/10.1126/science.1064088

[30]

Ma JH, Luan JW, Wang H, Wu PF, Ye XD, Wang Y, Ming AG, Liu SR. Nitrogen-fixing tree species rather than tree species diversity shape soil nematode communities in subtropical plantations. Geoderma, 2023, 436. ArticleID: 116561

[31]

Ma B, Sheng XJ, Zhou J, Nielsen UN, Wang XT, Ma MJ. Phosphorus addition ameliorates soil micro-food web simplification due to nitrogen enrichment but does not restore nematode community composition. Soil Biol Biochem, 2024, 195. ArticleID: 109447

[32]

Magneville C, Loiseau N, Albouy C, Casajus N, Claverie T, Escalas A, Leprieur F, Maire E, Mouillot D, Villéger S. mFD: an R package to compute and illustrate the multiple facets of functional diversity. Ecography, 2022, 2022(1. ArticleID: 05904

[33]

Mammola S, Carmona CP, Guillerme T, Cardoso P. Concepts and applications in functional diversity. Funct Ecol, 2021, 35(9): 1869-1885.

[34]

Mouillot D, Graham NAJ, Villéger S, Mason NWH, Bellwood DR. A functional approach reveals community responses to disturbances. Trends Ecol Evol, 2013, 28(3): 167-177.

[35]

Neher DA. Ecology of plant and free-living nematodes in natural and agricultural soil. Annu Rev Phytopathol, 2010, 48: 371-394.

[36]

Pang XY, Bao WK, Wu N. The effects of clear-felling subalpine coniferous forests on soil physical and chemical properties in the eastern Tibetan Plateau. Soil Use Manag, 2011, 27(2): 213-220.

[37]

Potapov AM, Beaulieu F, Birkhofer K, Bluhm SL, Degtyarev MI, Devetter M, Goncharov AA, Gongalsky KB, Klarner B, Korobushkin DI, Liebke DF, Maraun M, Mc Donnell RJ, Pollierer MM, Schaefer I, Shrubovych J, Semenyuk II, Sendra A, Tuma J, Tůmová M, Vassilieva AB, Chen TW, Geisen S, Schmidt O, Tiunov AV, Scheu S. Feeding habits and multifunctional classification of soil-associated consumers from protists to vertebrates. Biol Rev, 2022, 97(3): 1057-1117.

[38]

Pukkala T, Nuutinen Y, Muhonen T. A two-level optimization approach to tree-level planning in continuous cover forest management. J for Res, 2025, 36(1): 79.

[39]

Qi XX, Hu ZK, Li XP, Wang DY, Xu C, Hu F, Chen XY, Liu MQ. Land-use driven changes in elemental stoichiometry decouple the positive soil biodiversity-stability relationship. Soil Biol Biochem, 2025, 209. ArticleID: 109907

[40]

Qiang W, Gunina A, Kuzyakov Y, Luo RY, Zhang Y, Liu B, Pang XY. Shifts of understory vegetation induced by thinning drive the expansion of soil rare fungi. J Environ Manag, 2023, 342. ArticleID: 118119

[41]

Ren H, Corlett RT, Hui DF, Guo QF. Rewilding artificial forests in China and the world. The Innovation Life, 2025, 3(1. ArticleID: 100113

[42]

Ritter E, Bjørnlund L. Nitrogen availability and nematode populations in soil and litter after gap formation in a semi-natural beech-dominated forest. Appl Soil Ecol, 2005, 28(2): 175-189.

[43]

Schneider FD, Brose U, Rall BC, Guill C. Animal diversity and ecosystem functioning in dynamic food webs. Nat Commun, 2016, 7: 12718.

[44]

Schrader J, Craven D, Sattler C, Cámara-Leret R, Moeljono S, Kreft H. Life-history dimensions indicate non-random assembly processes in tropical island tree communities. Ecography, 2021, 44(3): 469-480.

[45]

Shen Y, Yang WQ, Zhang J, Xu ZF, Zhang L, Liu Y, Li H, You CM, Tan B. Forest gap size alters the functional diversity of soil nematode communities in Alpine forest ecosystems. Forests, 2019, 10(9. ArticleID: 806

[46]

Sun YX, Du XF, Li YB, Han X, Fang S, Geisen S, Li Q. Database and primer selections affect nematode community composition under different vegetations of Changbai Mountain. Soil Ecol Lett, 2023, 5(1): 142-150.

[47]

Tian AL, Halik Ü, Fu WT, Sawirdin S, Cheng SY, Lei JQ. Research history of forest gap as small-scale disturbances in forest ecosystems. Forests, 2024, 15(1): 21.

[48]

Tilman D. The ecological consequences of changes in biodiversity: a search for general principles. Ecology, 1999, 80(5): 1455-1474.

[49]

Tinya F, Csépányi P, Horváth CV, Kovács B, Németh C, Ódor P. Fine-scale interventions can reinforce the forest character of the understory vegetation–the effects of different artificial gaps in an oak-dominated forest. For Ecol Manag, 2025, 578. ArticleID: 122471

[50]

Tu CW, Lu QQ, Zhang Y, Tian JJ, Gao YR, Liu Y, Yang HB, Chen LH, Zhang J, Wang J, Xiao JJ. The soil nematode community indicates the soil ecological restoration of the Pinus massoniana plantation gap replanted with Cinnamomum longipaniculatum. Ecol Indic, 2022, 136. ArticleID: 108678

[51]

Tu CW, Zhang AJ, Luo RY, Qiang W, Zhang Y, Pang XY, Kuzyakov Y. Linking nematode trophic diversity to plantation identity and soil nutrient cycling. Geoderma, 2024, 448. ArticleID: 116945

[52]

Tu CW, Zhang AJ, Zhou Z, Zhang Y, Chen L, Potapov AM, Pang XY. Taxonomic assembly and trait partitioning contribute comparably to soil invertebrate functional diversity along secondary succession in high-elevation plateau ecosystems. Soil Biol Biochem, 2025, 211. ArticleID: 109995

[53]

Tu CW, Zhang Y, Dang XN, Liu Y, Huang YM, Liu GH, Hu ZD, Xiao JJ. Near-natural conversion of plantations primarily enhances the abundance and functionality of nematodes within medium-sized soil aggregates. J Environ Manag, 2025, 378. ArticleID: 124751

[54]

Tu CW, Zhang AJ, Zhang Y, Feng YL, Chen L, Pang XY. Soil fauna niche complementarity explains long-term soil multifunctionality differences between planted and natural subalpine forests. Geoderma, 2026, 465. ArticleID: 117653

[55]

van den Hoogen J, Geisen S, Routh D, Ferris H, Traunspurger W, Wardle DA, de Goede RGM, Adams BJ, Ahmad W, Andriuzzi WS, Bardgett RD, Bonkowski M, Campos-Herrera R, Cares JE, Caruso T, de Brito Caixeta L, Chen XY, Costa SR, Creamer R, Mauro da Cunha Castro J, Dam M, Djigal D, Escuer M, Griffiths BS, Gutiérrez C, Hohberg K, Kalinkina D, Kardol P, Kergunteuil A, Korthals G, Krashevska D, Kudrin AA, Li Q, Liang WJ, Magilton M, Marais M, Martín JAR, Matveeva E, Mayad EH, Mulder C, Mullin P, Neilson R, Duong Nguyen TA, Nielsen UN, Okada H, Rius JEP, Pan KW, Peneva V, Pellissier L, Carlos Pereira da Silva J, Pitteloud P, Powers TO, Powers K, Quist CW, Rasmann S, Moreno SS, Scheu S, Setälä H, Sushchuk A, Tiunov AV, Trap J, van der Putten W, Vestergård M, Villenave C, Waeyenberge A, Wall DH, Wilschut R, Wright DG, Yang JI, Crowther TW. Soil nematode abundance and functional group composition at a global scale. Nature, 2019, 572(7768): 194-198.

[56]

Wang JQ, Shi XZ, Lucas-Borja ME, Guo QL, Mao JY, Tan YY, Zhang GY (2023) Soil nematode abundances drive agroecosystem multifunctionality under short-term elevated CO2 and O3. Glob Change Biol 29(6):1618–1627. https://doi.org/10.1111/gcb.16546

[57]

Xing W, Hu N, Li ZF, Feng LS, Zhang WD, Du Preez G, Zhang HM, Li DC, Lu SB, Chang SX, Zhang QW, Lou YL. Soil enzyme profile analysis for indicating decomposer micro-food web. iMeta, 2024, 3(1. ArticleID: e161

[58]

Xu JX, Lie GW, Xue L. Effects of gap size on diversity of soil fauna in a Cunninghamia lanceolata stand damaged by an ice storm in southern China. J for Res, 2016, 27(6): 1427-1434.

[59]

Yang B, Pang XY, Bao WK, Zhou KX. Thinning-induced canopy opening exerted a specific effect on soil nematode community. Ecol Evol, 2018, 8(8): 3851-3861.

[60]

Yang HJ, Pan C, Wu Y, Qing SQ, Wang ZB, Wang DH. Response of understory plant species richness and tree regeneration to thinning in Pinus tabuliformis plantations in northern China. For Ecosyst, 2023, 10. ArticleID: 100105

[61]

Yeates GW. Abundance, diversity, and resilience of nematode assemblages in forest. Can J for Res, 2007, 37: 216-225.

[62]

Yin HF, Su Y, Zeng J, Li XW, Fan C, Lu JZ, Zhou Z, Yu AW, Wang SM, Scheu S, Krashevska V. Forest gap regulates soil nematode community through understory plant diversity and soil pH. Geoderma, 2024, 451. ArticleID: 117086

[63]

Zhang Y, Zhang DJ, Li X, Zhang J. Contribution of soil fauna to the degradation of recalcitrant components in Cinnamomum camphora foliar litter in different-sized gaps in Pinus massoniana plantations. J for Res, 2019, 30(3): 931-941.

[64]

Zhang ZW, Li Q, Hu YY, Wei HW, Hou SL, Yin JX, Lü XT (2022) Nitrogen and phosphorus additions interactively affected composition and carbon budget of soil nematode community in a temperate steppe. Plant Soil 473(1/2):109–121. https://doi.org/10.1007/s11104-021-05145-y

[65]

Zhang AN, Chen SY, Chen JW, Cui HW, Jiang XX, Xiao S, Wang JJ, Gao HN, An LZ, Cardoso P. Shrub and precipitation interactions shape functional diversity of nematode communities on the Qinghai-Tibet Plateau. Glob Change Biol, 2023, 29(10): 2746-2758.

[66]

Zhang CZ, Wright IJ, Nielsen UN, Geisen S, Liu MQ. Linking nematodes and ecosystem function: a trait-based framework. Trends Ecol Evol, 2024, 39(7): 644-653.

[67]

Zhang Y, Zhou Z, Junggebauer A, Pollierer MM, Scheu S. Effects of tree fall on soil Collembola: disentangling the role of gap formation and deadwood addition. Geoderma, 2025, 455. ArticleID: 117217

[68]

Zhang Y, Junggebauer A, Pollierer MM, Scheu S, Zhou Z. Contrasting effects of deadwood and gaps on the trophic structure of forest soil microarthropods. Funct Ecol, 2026, 40(1): 150-162.

[69]

Zhu BJ, Whalen JK, Wu JT, Yang JN, Mao XR, Wan BB, Tian SY, Hu F, Chen XY, Liu MQ. Soil food web structure coordinated by soil omnivores sustains soil multifunctionality in moderate vermicompost amended fields. Soil Biol Biochem, 2024, 192. ArticleID: 109391

RIGHTS & PERMISSIONS

Northeast Forestry University

PDF

0

Accesses

0

Citation

Detail

Sections
Recommended

/