Long-term passive rewilding enhances total carbon stock but reduces annual productivity in abandoned bamboo forests

Abstract

Forest passive rewilding, a nature-based restoration approach that minimizes human disturbances, supports forest succession and ecosystem functional restoration. However, its effect on ecosystem carbon stock and forest productivity remains elusive and context-dependent. This study investigated the long-term (up to 30 years) effects of rewilding on forest structure, above- and below-ground carbon pools, and annual productivity in a subtropical forest dominated by the fast-growing species, Moso bamboo (Phyllostachys edulis). Compared to conventional management, rewilding significantly restructured forests by increasing stand density and the proportion of old bamboo culms (≥ 7 years old), and decreasing understory plant richness. However, the response of bamboo growth varied across rewilding durations and culm ages. Only long-term rewilding (30 years) improved the growth of young bamboo culms (≤ 5 years old, increased by 12–35%, P < 0.05) through soil improvement. These changes led to increased total bamboo biomass after rewilding. Moreover, compared to regular harvest in conventional bamboo forest, rewilding retained carbon from harvested biomass to in-situ (standing bamboo, deadwood and soil) carbon stocks. This shift in carbon allocation was associated with a reduction in annual gross primary productivity (AGPP, – 61%, P < 0.001) and an increase in total soil organic carbon stock (+ 63%, P < 0.05). Ultimately, the structural equation model indicated that ecosystem productivity and carbon stock were mainly influenced by the bamboo and its stand structure, as mediated by rewilding duration, rather than understory plants. Together, these findings provide an integrative framework for evaluating carbon sequestration strategy after passive rewilding, and offer critical insights for understanding sustainable forest management strategies that balance short-term productivity with long-term carbon storage to support climate-mitigation goals.

Keywords

Carbon sequestration / Annual productivity / Forest rewilding / Plant diversity / Moso bamboo

References

[1] Atkinson J, Brudvig LA, Mallen-Cooper M, Nakagawa S, Moles AT, Bonser SP. Terrestrial ecosystem restoration increases biodiversity and reduces its variability, but not to reference levels: a global meta-analysis. Ecol Lett, 2022, 25(7): 1725-1737.

[2] Bowd EJ, Banks SC, Strong CL, Lindenmayer DB. Long-term impacts of wildfire and logging on forest soils. Nat Geosci, 2019, 12(2): 113-118.

[3] Brancalion PHS, Hua FY, Joyce FH, Antonelli A, Holl KD. Moving biodiversity from an afterthought to a key outcome of forest restoration. Nat Rev Biodivers, 2025, 1(4): 248-261.

[4] Bray RH, Kurtz LT. Determination of total, organic, and available forms of phosphorus in soils. Soil Sci, 1945, 59(1): 39-46.

[5] Chen SP, Wang WT, Xu WT, Wang Y, Wan HW, Chen DM, Tang ZY, Tang XL, Zhou GY, Xie ZQ, Zhou DW, Shangguan Z, Huang JH, He JS, Wang YF, Sheng JD, Tang LS, Li XR, Dong M, Wu Y, Wang QF, Wang ZH, Wu JG, Chapin FSIII, Bai YF. Plant diversity enhances productivity and soil carbon storage. Proc Natl Acad Sci U S A, 2018, 115(16): 4027-4032.

[6] Chen X, Chen X, Huang SQ, Fang DM. Impacts of Moso bamboo (Phyllostachys pubescens) invasion on species diversity and aboveground biomass of secondary coniferous and broad-leaved mixed forest. Front Plant Sci, 2022, 13. ArticleID: 1001785

[7] Chen XL, Taylor AR, Reich PB, Hisano M, Chen HYH, Chang SX. Tree diversity increases decadal forest soil carbon and nitrogen accrual. Nature, 2023, 618(7963): 94-101.

[8] Chen C, Xiao WY, Chen HYH. Meta-analysis reveals global variations in plant diversity effects on productivity. Nature, 2025, 638(8050): 435-440.

[9] Constenla-Villoslada S, Liu YY, Wen JM, Sun Y, Chonabayashi S. Large-scale land restoration improved drought resilience in Ethiopia’s degraded watersheds. Nat Sustain, 2022, 5(6): 488-497.

[10] Crouzeilles R, Curran M, Ferreira MS, Lindenmayer DB, Grelle CEV, Rey Benayas JM. A global meta-analysis on the ecological drivers of forest restoration success. Nat Commun, 2016, 7. ArticleID: 11666

[11] Dai W, Zhao KL, Gao ZQ, Liu KH, Zhang F, Fu WJ. Spatial variation characteristics of carbon density and storage in forest ecosystems in a typical subtropical region. Acta Ecol Sin, 2017, 37(22): 7528-7538.

[12] Dong HY, Ge JF, Sun K, Wang BZ, Xue JM, Wakelin SA, Wu JS, Sheng WX, Liang CF, Xu QF, Jiang PK, Chen JH, Qin H. Change in root-associated fungal communities affects soil enzymatic activities during Pinus massoniana forest development in subtropical China. For Ecol Manag, 2021, 482. ArticleID: 118817

[13] Fang QY, Zhou S, Hu S, Ye CY, Fang DM, Mei TT, Zhou GM. Above-ground biomass of Moso bamboo forests in China influenced by climate, soil and topography: a meta-analysis. Adv Bamboo Sci, 2024, 8. ArticleID: 100091

[14] Gamfeldt L, Snäll T, Bagchi R, Jonsson M, Gustafsson L, Kjellander P, Ruiz-Jaen MC, Fröberg M, Stendahl J, Philipson CD, Mikusiński G, Andersson E, Westerlund B, Andrén H, Moberg F, Moen J, Bengtsson J. Higher levels of multiple ecosystem services are found in forests with more tree species. Nat Commun, 2013, 4. ArticleID: 1340

[15] Ge ZW, Yu T, Tian XY, Chen XX, Yao YW, Mao LF. Analysis of understory plant community assembly differences in moso bamboo forests in the subtropical evergreen broad-leaved forest region of Eastern China. Forests, 2025, 16(3): 478.

[16] Hartley IP, Garnett MH, Sommerkorn M, Hopkins DW, Fletcher BJ, Sloan VL, Phoenix GK, Wookey PA. A potential loss of carbon associated with greater plant growth in the European Arctic. Nat Clim Change, 2012, 2(12): 875-879.

[17] Hua FY, Bruijnzeel LA, Meli P, Martin PA, Zhang J, Nakagawa S, Miao XR, Wang WY, McEvoy C, Peña-Arancibia JL, Brancalion PHS, Smith P, Edwards DP, Balmford A. The biodiversity and ecosystem service contributions and trade-offs of forest restoration approaches. Science, 2022, 376(6595): 839-844.

[18] Larpkern P, Moe SR, Totland Ø. Bamboo dominance reduces tree regeneration in a disturbed tropical forest. Oecologia, 2011, 165(1): 161-168.

[19] Lei GS, Han ZX, Wang XY, Malacrinò A, Kang T, Zhang DD, Zhang JY, Zhang Z, Wu HM. Synthetic microbial communities rescues strawberry from soil-borne disease by enhancing soil functional microbial abundance and multifunctionality. J Adv Res, 2026, 83: 113-128.

[20] Li Q, Ma QH, Gao JP, Zhang JB, Li YF, Shi M, Peng CH, Song XZ. Stumps increased soil respiration in a subtropical Moso bamboo (Phyllostachys edulis) plantation under nitrogen addition. Agric for Meteor, 2022, 323. ArticleID: 109047

[21] Li JJ, Huang YM, Chen LH, Gao S, Zhang J, Zhang DJ. Understory plant diversity and phenolic allelochemicals across a range of Eucalyptus grandis plantation ages. J for Res, 2023, 34: 1577-1590.

[22] Li Q, Zhang C, Shi M, Lv JH, Peng CH, Zhang JB, Chang SX, Cao TT, Li T, Song XZ. Long-term nitrogen addition has a positive legacy effect on soil respiration in subtropical Moso bamboo forests. Geoderma, 2024, 452. ArticleID: 117092

[23] Li SB, Zhao CS, Ren WJ, Zhang Z, Wang W, Zhang Q, Ma MY, Zhao GX, Chen FY, Liu H, Peng WY, Wang ZH, Jiang W, Wang LL, Zhang WP, Zhang Y, Ma JR, Yu WJ, Wang ZK, Gao YZ, Chen JL, Zhang XB, Zhou DF, Lv J, Wang S, Casals Mercadal E, Peng HD, Zhao WJ, Zeng ML. The interplay of environmental and human microbiomes: a review of diversity, functions, and implications for one health. Innovation Life, 2026, 4. ArticleID: 100208

[24] Li YY, Wang N, Latiff ARA. Development of the bamboo forest economy: reviewing China’s 'bamboo as a substitute for plastic initiative' and its development. Adv Bamboo Sci, 2025, 11. ArticleID: 100130

[25] Lu ZJ, Franklin SB. Clonal integration and regeneration in bamboo Bashania fargesii. For Ecol Manag, 2022, 523. ArticleID: 120504

[26] Luo WX, Ma YX, Luo J, Han SX, Sun CX, You ZY, Du HH, Li ZF, Liu JC, Tao JP. Understory bamboo removal impacts on woody seedling regeneration in forest ecosystems: a meta-analysis. Ecol Process, 2024, 13(1. ArticleID: 47

[27] Luo YH, Cheng JM, Mahmood M, Wei SH, Peng LQ, Zhang C, Zhang M, Zhang JF, yang H, Wang JZ, Hao ZG, Yan XF. Plant community restoration time is a key factor affecting plant nutrient and species richness. Sci Rep, 2025, 15. ArticleID: 15752

[28] Lv JH, Li Q, Cao TT, Shi M, Peng CH, Deng L, Song XZ. A compartmentation approach to deconstruct ecosystem carbon fluxes of a Moso bamboo forest in subtropical China. For Ecosyst, 2025, 12. ArticleID: 100286

[29] Makelele IA, Verheyen K, Boeckx P, Cizungu Ntaboba L, Mujinya Bazirake B, Ewango C, Bauters M. Afrotropical secondary forests exhibit fast diversity and functional recovery, but slow compositional and carbon recovery after shifting cultivation. J Veg Sci, 2021, 32(5. ArticleID: e13071

[30] Makita A. Density regulation during the regeneration of two monocarpic bamboos: self-thinning or intraclonal regulation?. J Veg Sci, 1996, 7(2): 281-288.

[31] Markuljaková K, Mikoláš M, Svitok M, Meigs GW, Keeton WS, Kozák D, Pavlin J, Gloor R, Kalaš M, Ferenčík M, Ralhan D, Frankovič M, Hofmeister J, Dúhová D, Mejstrík M, Dušátko M, Veber A, Knír T, Svoboda M. Rewilding beech-dominated temperate forest ecosystems: effects on carbon stocks and biodiversity indicators. iForest, 2025, 18(1): 1-9.

[32] Ortega-Barrueta U, Sertutxa U, Ametzaga-Arregi I, Montaño-Lekue M, Peña L. Multitaxon biodiversity and functional recovery in restored native forests in a biosphere reserve. For Ecol Manag, 2025, 596. ArticleID: 123057

[33] Pan CY, Wang GY, Xu L, Li C, Shrestha A, Ying MJ, Lu WM, Innes JL, Kozak R, Zhou GM. Bamboo forests in Anji, China: an emerging nature-based solution to tackle climate change. Adv Bamboo Sci, 2025, 10. ArticleID: 100126

[34] Perino A, Pereira HM, Navarro LM, Fernández N, Bullock JM, Ceaușu S, Cortés-Avizanda A, van Klink R, Kuemmerle T, Lomba A, Pe’er G, Plieninger T, Rey Benayas JM, Sandom CJ, Svenning JC, Wheeler HC. Rewilding complex ecosystems. Science, 2019, 364(6438. ArticleID: eaav5570

[35] Pohlman CL, Tng DYP, Florentine SK. Do primary rainforest tree species recruit into passively and actively restored tropical rainforest?. For Ecol Manage, 2021, 496. ArticleID: 119453

[36] Qian F, Song HY, Chen M, Zeng JQ, Dang CQ, Tao JP. Multivariate path analysis of the relationships between seedling regeneration and environmental factors beneath a dwarf bamboo understory. Ecol Evol, 2019, 9(18): 10277-10290.

[37] Qin FC, Liu S, Yu SX. Effects of allelopathy and competition for water and nutrients on survival and growth of tree species in Eucalyptus urophylla plantations. For Ecol Manage, 2018, 424: 387-395.

[38] Raaijmakers JM, Kiers ET. Rewilding plant microbiomes. Science, 2022, 378(6620): 599-600.

[39] Rastetter EB, Kling GW, Shaver GR, Crump BC, Gough L, Griffin KL. Ecosystem recovery from disturbance is constrained by N cycle openness, vegetation-soil N distribution, form of N losses, and the balance between vegetation and soil-microbial processes. Ecosystems, 2021, 24(3): 667-685.

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

[41] Shi M, Zhang JB, Sun JL, Li Q, Lin XC, Song XZ. Unequal nitrogen translocation pattern caused by clonal integration between connected ramets ensures necessary nitrogen supply for young Moso bamboo growth. Environ Exp Bot, 2022, 200. ArticleID: 104900

[42] Song XZ, Zhou GM, Jiang H, Yu SQ, Fu JH, Li WZ, Wang WF, Ma ZH, Peng CH. Carbon sequestration by Chinese bamboo forests and their ecological benefits: assessment of potential, problems, and future challenges. Environ Rev, 2011, 19: 418-428.

[43] Song XZ, Peng CH, Ciais P, Li Q, Xiang WH, Xiao WF, Zhou GM, Deng L. Nitrogen addition increased CO₂ uptake more than non-CO₂ greenhouse gases emissions in a Moso bamboo forest. Sci Adv, 2020, 6(12. ArticleID: eaaw5790

[44] Spake R, Ezard THG, Martin PA, Newton AC, Doncaster CP. A meta-analysis of functional group responses to forest recovery outside of the tropics. Conserv Biol, 2015, 29(6): 1695-1703.

[45] Su YX, Zhang CQ, Cescatti A, Yu KL, Ciais P, Smith T, Shang JL, Carnicer J, Liu J, Chen JM, Green JK, Wu JP, Ponce-Campos GE, Zhang YG, Zuo ZY, Liao JB, Wu JP, Lafortezza R, Yan K, Yang XQ, Liu LY, Ren JS, Yuan WP, Chen XZ, Wu CY, Zhou WQ. Pervasive but biome-dependent relationship between fragmentation and resilience in forests. Nat Ecol Evol, 2025, 9(9): 1670-1684.

[46] Teng QM, Lu XN, Zhang QQ, Cai LL, Sardar MF, Li YF, Abbas T, Li Y, Chang SX, Li YC. Litterfall quality modulates soil ammonium and nitrate supply through altering microbial function in bamboo encroachment of broadleaf forests. Geoderma, 2023, 437. ArticleID: 116592

[47] Token S, Jiang LM, Zhang L, Lv GH. Effects of plant diversity on primary productivity and community stability along soil water and salinity gradients. Glob Ecol Conserv, 2022, 36. ArticleID: e02095

[48] Ullah S, Wu JP, Shah JA, Wang XM, Lyu YM, Guo ZW, Ali K, Chen DY, Han S. Tree diversity drives understory carbon storage rather than overstory carbon storage across forest types. J for Res, 2024, 35: 125.

[49] Wang T, Dong LB, Liu ZG. Factors driving native tree species restoration in plantations and tree structure conversion in Chinese temperate forests. For Ecol Manag, 2022, 507. ArticleID: 119989

[50] Wang SL, Wang XJ, Cao WX. Fertile island soils promote the restoration of shrub patches in burned areas in arid saline land. Fire, 2023, 6(9): 341.

[51] Wang YJ, Lin YB, Zhang L, Liu SY, Wang J, Tian Y, Campbell DE, Lin RY, Ren H, Lu HF. Long-term effects of intercropping on multi-trophic structure and bio-thermodynamic health of mixed Eucalyptus-native tree plantations. J Appl Ecol, 2024, 61(1): 103-119.

[52] Wang LH, Wei FL, Tagesson T, Fang ZX, Svenning JC. Transforming forest management through rewilding: enhancing biodiversity, resilience, and biosphere sustainability under global change. One Earth, 2025, 8(3. ArticleID: 101195

[53] Wang ZK, Li Q, Shi M, Leite MFA, Chen XL, Kuramae EE, Cordovez V, Cao TT, Zhu CL, Zhou LB, Yu WJ, Tang ZY, Peng CH, Song XZ. Compartmentalized homeostasis drives high bamboo forest productivity under nutrient imbalance. Adv Sci, 2026, 13(11. ArticleID: e17442

[54] Wei X, Wei SP, Hao DN, Jia L, Liang WJ. Optimizing adaptive disturbance in planted forests: resource allocation strategies for sustainable regeneration from seedlings to saplings. Plant Cell Environ, 2025, 48(11): 8114-8126.

[55] Wills J, Herbohn J, Wells J, Maranguit Moreno MO, Ferraren A, Firn J. Seedling diversity in actively and passively restored tropical forest understories. Ecol Appl, 2021, 31(3. ArticleID: e02286

[56] Woś B, Chodak M, Likus-Cieślik J, Misebo AM, Pietrzykowski M. Regeneration of post-mining and post-fire soil function by assessment of tree nutrient status: evidence from pioneer and N-fixing species. Land Degrad Dev, 2024, 35(15): 4521-4534.

[57] Wu CY, Deng L, Huang CB, Chen YF, Peng CH. Effects of vegetation restoration on soil nutrients, plant diversity, and its spatiotemporal heterogeneity in a desert–oasis ecotone. Land Degrad Dev, 2021, 32(2): 670-683.

[58] Xie YY, Zhang W, Guo ZW, Du XH, Fan LL, Chen SL, Dong YW. Effects of vegetation succession on soil microbial stoichiometry in Phyllstachys edulis stands following abandonment. Sci Total Environ, 2023, 895. ArticleID: 164971

[59] Xiong L, Yuan CX, Wu QQ, Fornara DA, Heděnec P, Chen SY, Peng Y, Zhao ZM, Wu FZ, Yue K. Understory vegetation removal significantly affected soil biogeochemical properties in forest ecosystems. Appl Soil Ecol, 2024, 193. ArticleID: 105132

[60] Yuen JQ, Fung T, Ziegler AD. Carbon stocks in bamboo ecosystems worldwide: estimates and uncertainties. For Ecol Manag, 2017, 393: 113-138.

[61] Zhai KY, Yin P, Revillini D, Chen XL, Liu SG, Lou YL, Zhang WD, Wang SL, Zhou GY, Delgado-Baquerizo M. Subordinate tree species diversity explains carbon stock better than dominant tree diversity after decades of forest rewilding. J Appl Ecol, 2025, 62(3): 604-616.

[62] Zhang JB, Li Q, Lv JH, Peng CH, Gu ZK, Qi LH, Song XZ, Song XZ. Management scheme influence and nitrogen addition effects on soil CO₂, CH₄, and N₂O fluxes in a Moso bamboo plantation. For Ecosyst, 2021, 8: 6.

[63] Zhang HL, Liu SR, Yu JY, Li JW, Shangguan ZP, Deng L. Thinning increases forest ecosystem carbon stocks. For Ecol Manag, 2024, 555. ArticleID: 121702

[64] Zhang L, Liu SY, Wang YJ, Zhang FW, Luo QY, Lin YB, Hui DF, Fu SL, Wang J, Liu ZF, Lin RY, Ren H, Lu HF. Optimizing multiple ecosystem services in Eucalyptus plantations: dual effects of intercropping and restoration. J Clean Prod, 2024, 483. ArticleID: 144288

[65] Zhang WY, Shi M, Yang KB, Zhang JB, Gao ZM, El-Kassaby YA, Li Q, Cao TT, Deng SX, Qing HS, Wang ZK, Song XZ. Regulatory networks of senescence-associated gene-transcription factors promote degradation in Moso bamboo shoots. Plant Cell Environ, 2024, 47(9): 3654-3667.

[66] Zhang JB, Shi M, Zhu CL, Yang KB, Li Q, Song XM, Gao ZM, Cao TT, Zhu DZ, Song XZ. Stable isotope labelling and gene expression analysis reveal dynamic nitrogen-supply mechanisms for rapid growth of Moso bamboo. Hortic Res, 2025, 12(6. ArticleID: uhaf062

[67] Zheng AY, Lv JH. Spatial patterns of bamboo expansion across scales: how does Moso bamboo interact with competing trees?. Landsc Ecol, 2023, 38(12): 3925-3943.

[68] Zhou GY, Lucas-Borja ME, Liu SG, Hu HW, He JZ, Wang XX, Jiang Z, Zhou XH, Delgado-Baquerizo M. Plant and soil biodiversity is essential for supporting highly multifunctional forests during Mediterranean rewilding. Funct Ecol, 2023, 37(2): 420-431.

[69] Zhou J, Shao GD, Liu EK, Liu Q, Yan CR, Alharbi SA, Filimonenko E, Mei XR, Kuzyakov Y. Climate warming and agronomic practice interactively alter soil carbon stock in dry farmland in China. Commun Earth Environ, 2025, 6: 788.

RIGHTS & PERMISSIONS

Northeast Forestry University