Abstract
Plant aboveground biomass reflects the resilience of ecosystem productivity to different environmental factors and plays an important role in biodiversity and ecosystem carbon cycles. Although many studies have investigated the relationship between aboveground biomass and species diversity, stand structural diversity is an important factor affecting ecosystem function. The mechanism of how climate and soil factors affect aboveground biomass through species diversity and structural diversity remains unclear. Here, combing data from 8 locations of Abies fargesii var. faxoniana primary forest, a structural equation model was used to study the effects of climate factors, soil factors, species diversity, and structural diversity on aboveground biomass. Our results demonstrated that climate factors affect aboveground biomass mainly through the indirect effect of species diversity. Species diversity indirectly affects aboveground biomass through its effects on structural diversity. Both direct effects of soil factors and indirect effects through structural diversity can significantly affect aboveground biomass. Therefore, maintaining higher diversity of stand structure and soil nutrients can improve aboveground biomass, and thus better improve ecosystem functions such as carbon storage.
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The data that support the findings of this study are available for reasonable request from the corresponding author.
References
Adler PB, Seabloom EW, Borer ET et al (2011) Productivity is a poor predictor of plant species richness. Science 333:1750–1753. https://doi.org/10.1126/science.1204498
Ali A, Yan ER (2017a) The forest strata-dependent relationship between biodiversity and aboveground biomass within a subtropical forest. For Ecol Manage 401:125–134. https://doi.org/10.1016/j.foreco.2017.06.056
Ali A, Yan ER (2017b) Functional identity of overstorey tree height and understorey conservative traits drive aboveground biomass in a subtropical forest. Ecol Ind 83:158–168. https://doi.org/10.1016/j.ecolind.2017.07.054
Ali A, Yan ER (2018) The mediation roles of intraspecific and interspecific functional trait diversity for linking the response of aboveground biomass to species richness across forest strata in a subtropical forest. Ecol Ind 85:493–501. https://doi.org/10.1016/j.ecolind.2017.10.057
Ali A, Yan ER, Chen HYH et al (2016) Stand structural diversity rather than species diversity enhances aboveground carbon storage in secondary subtropical forests in Eastern China. Biogeosciences 13:4627–4635. https://doi.org/10.5194/bg-13-4627-2016
Ali A, Chen HYH, You WH et al (2019a) Multiple abiotic and biotic drivers of aboveground biomass shift with forest stratum. For Ecol Manage 436:1–10. https://doi.org/10.1016/j.foreco.2019.01.007
Ali A, Lin SL, He JK et al (2019b) Big-sized trees overrule remaining trees’ attributes and species richness as determinants of aboveground biomass in tropical forests. Glob Change Biol 25:2810–2824. https://doi.org/10.1111/gcb.14707
Ali A, Sanaei A, Li M et al (2020) Impacts of climatic and edaphic factors on the diversity, structure and biomass of species-poor and structurally-complex forests. Sci Total Environ 706:135719. https://doi.org/10.1016/j.scitotenv.2019.135719
Anderson RC, Loucks OL, Swain AM (1969) Herbaceous response to canopy cover, light intensity, and throughfall precipitation in coniferous forests. Ecology 50:255–263. https://doi.org/10.2307/1934853
Antonelli A, Nylander JAA, Persson C et al (2009) Tracing the impact of the Andean uplift on Neotropical plant evolution. Proc Natl Acad Sci 106:9749–9754. https://doi.org/10.1073/pnas.0811421106
Bar-On YM, Phillips R, Milo R (2018) The biomass distribution on Earth. Proc Natl Acad Sci 115:6506–6511. https://doi.org/10.1073/pnas.1711842115
Cao XW, Shi ZM, Chen J et al (2022) Extracellular enzyme characteristics and microbial metabolic limitation in soil of subalpine forest ecosystems on the eastern Qinghai-Tibetan Plateau. Plant Soil 479:337–353. https://doi.org/10.1007/s11104-022-05521-2
Cardinale BJ, Wright JP, Cadotte MW et al (2007) Impacts of plant diversity on biomass production increase through time because of species complementarity. Proc Natl Acad Sci 104:18123–18128. https://doi.org/10.1073/pnas.0709069104
Chen J, Shi Z, Liu S et al (2022) Altitudinal variation influences soil fungal community composition and diversity in Alpine-Gorge region on the eastern Qinghai-Tibetan Plateau. Journal of Fungi 8:807. https://doi.org/10.3390/jof8080807
Chen M, Shi Z, Liu S et al (2023) Leaf functional traits have more contributions than climate to the variations of leaf stable carbon isotope of different plant functional types on the eastern Qinghai-Tibetan Plateau. Sci Total Environ 871:162036. https://doi.org/10.1016/j.scitotenv.2023.162036
Craine JM, Dybzinski R, Robinson D (2013) Mechanisms of plant competition for nutrients, water and light. Funct Ecol 27:833–840. https://doi.org/10.1111/1365-2435.12081
Dănescu A, Albrecht AT, Bauhus J (2016) Structural diversity promotes productivity of mixed, uneven-aged forests in southwestern Germany. Oecologia 182:319–333. https://doi.org/10.1007/s00442-016-3623-4
Ding Y, Zang R (2021) Determinants of aboveground biomass in forests across three climatic zones in China. For Ecol Manage 482:118805. https://doi.org/10.1016/j.foreco.2020.118805
Dormann CF, McPherson JM, Araújo MB et al (2007) Methods to account for spatial autocorrelation in the analysis of species distributional data: a review. Ecography 30:609–628. https://doi.org/10.1111/j.2007.0906-7590.05171.x
Fotis AT, Murphy SJ, Ricart RD et al (2018) Above-ground biomass is driven by mass-ratio effects and stand structural attributes in a temperate deciduous forest. J Ecol 106:561–570. https://doi.org/10.1111/1365-2745.12847
Fraser LH, Pither J, Jentsch A et al (2015) Worldwide evidence of a unimodal relationship between productivity and plant species richness. Science 349:302–305. https://doi.org/10.1126/science.aab3916
Fu X, Yang F, Wang J et al (2015) Understory vegetation leads to changes in soil acidity and in microbial communities 27 years after reforestation. Sci Total Environ 502:280–286. https://doi.org/10.1016/j.scitotenv.2014.09.018
Gao Z, Wang Q, Hu Z et al (2019) Comparing independent climate-sensitive models of aboveground biomass and diameter growth with their compatible simultaneous model system for three larch species in China. Int J Biomath 12:1950053. https://doi.org/10.1142/S1793524519500530
Garnier E, Cortez J, Billès G et al (2004) Plant functional markers capture ecosystem properties during secondary succession. Ecology 85:2630–2637. https://doi.org/10.1890/03-0799
Grace JB, Anderson TM, Seabloom EW et al (2016) Integrative modelling reveals mechanisms linking productivity and plant species richness. Nature 529:390–393. https://doi.org/10.1038/nature16524
Grace JB (2006) Structural equation modeling and natural systems. Cambridge University Press, pp 1–378
Grime JP (1998) Benefits of plant diversity to ecosystems: immediate, filter and founder effects. J Ecol 86:902–910. https://doi.org/10.1046/j.1365-2745.1998.00306.x
Gröm** U (2007) Relative importance for linear regression in R: the package relaimpo. J Stat Softw 17:1–27. https://doi.org/10.18637/jss.v017.i01
Hutchinson MF, Xu TB (2013) ANUSPLIN version 4.4 user guide. Australian National University, Canberra
Jimenez-Alfaro B, Girardello M, Chytry M et al (2018) History and environment shape species pools and community diversity in European beech forests. Nat Ecol Evolut 2:483–490. https://doi.org/10.1038/s41559-017-0462-6
** Y, Liu C, Qian SS et al (2022) Large-scale patterns of understory biomass and its allocation across China’s forests. Sci Total Environ 804:150169. https://doi.org/10.1016/j.scitotenv.2021.150169
Li W, Li J, Liu S et al (2017) Magnitude of species diversity effect on aboveground plant biomass increases through successional time of abandoned farmlands on the eastern Tibetan Plateau of China. Land Degrad Dev 28:370–378. https://doi.org/10.1002/ldr.2607
Li F, Shi Z, Liu S et al (2023) Soil properties and plant diversity co-regulate ecosystem multifunctionality of subalpine primary dark coniferous forest on the eastern Qinghai-Tibetan Plateau. Plant Soil 493:207–219. https://doi.org/10.1007/s11104-023-06222-0
Liang J, Crowther TW, Picard N et al (2016) Positive biodiversity-productivity relationship predominant in global forests. Science 354:aaf8957. https://doi.org/10.1126/science.aaf8957
Lie Z, Lin W, Huang W et al (2019) Warming changes soil N and P supplies in model tropical forests. Biol Fertil Soils 55:751–763. https://doi.org/10.1007/s00374-019-01382-7
Liu Y, Su X, Shrestha N et al (2019) Effects of contemporary environment and Quaternary climate change on drylands plant diversity differ between growth forms. Ecography 42:334–345. https://doi.org/10.1111/ecog.03698
Liu S, Luo D, Cheng R et al (2021) Temporal variability in soil net nitrogen mineralization among forest regeneration patterns in eastern Tibetan Plateau. Ecol Ind 128:107811. https://doi.org/10.1016/j.ecolind.2021.107811
Liu S, Xu G, Chen H et al (2023) Contrasting responses of soil microbial biomass and extracellular enzyme activity along an elevation gradient on the eastern Qinghai-Tibetan Plateau. Front Microbiol 14:974316. https://doi.org/10.3389/fmicb.2023.974316
Loreau M, Hector A (2001) Partitioning selection and complementarity in biodiversity experiments. Nature 412:72–76. https://doi.org/10.1038/35083573
Luo YH, Cadotte MW, Burgess KS et al (2019) Greater than the sum of the parts: how the species composition in different forest strata influence ecosystem function. Ecol Lett 22:1449–1461. https://doi.org/10.1111/ele.13330
Luo Y, Wang X, Ouyang Z et al (2020) A review of biomass equations for China’s tree species. Earth Syst Sci Data 12:21–40. https://doi.org/10.5194/essd-12-21-2020
Luo Y, Zhao X, Li Y et al (2021) Wind disturbance on litter production affects soil carbon accumulation in degraded sandy grasslands in semi-arid sandy grassland. Ecol Eng 171:106373. https://doi.org/10.1016/j.ecoleng.2021.106373
Maestre FT, Castillo-Monroy AP, Bowker MA et al (2012) Species richness effects on ecosystem multifunctionality depend on evenness, composition and spatial pattern. J Ecol 100:317–330. https://doi.org/10.1111/j.1365-2745.2011.01918.x
Michaletz ST, Kerkhoff AJ, Enquist BJ (2018) Drivers of terrestrial plant production across broad geographical gradients. Glob Ecol Biogeogr 27:166–174. https://doi.org/10.1111/geb.12685
Ouyang S, **ang W, Wang X et al (2016) Significant effects of biodiversity on forest biomass during the succession of subtropical forest in south China. For Ecol Manage 372:291–302. https://doi.org/10.1016/j.foreco.2016.04.020
Palpurina S, Wagner V, Von Wehrden H et al (2017) The relationship between plant species richness and soil pH vanishes with increasing aridity across Eurasian dry grasslands. Glob Ecol Biogeogr 26:425–434. https://doi.org/10.1111/geb.12549
Paoli GD, Curran LM, Zak DR (2005) Phosphorus efficiency of Bornean rain forest productivity: evidence against the unimodal efficiency hypothesis. Ecology 86:1548–1561. https://doi.org/10.1890/04-1126
Poorter L, van der Sande MT, Thompson J et al (2015) Diversity enhances carbon storage in tropical forests. Glob Ecol Biogeogr 24:1314–1328. https://doi.org/10.1111/geb.12364
Potter KM, Woodall CW (2014) Does biodiversity make a difference? Relationships between species richness, evolutionary diversity, and aboveground live tree biomass across U.S. forests. For Ecol Manage 321:117–129. https://doi.org/10.1016/j.foreco.2013.06.026
Quesada CA, Phillips OL, Schwarz M et al (2012) Basin-wide variations in Amazon forest structure and function are mediated by both soils and climate. Biogeosciences 9:2203–2246. https://doi.org/10.5194/bg-9-2203-2012
Rüger N, Wirth C, Wright SJ et al (2012) Functional traits explain light and size response of growth rates in tropical tree species. Ecology 93:2626–2636. https://doi.org/10.1890/12-0622.1
Schnabel F, Schwarz JA, Dănescu A et al (2019) Drivers of productivity and its temporal stability in a tropical tree diversity experiment. Glob Change Biol 25:4257–4272. https://doi.org/10.1111/gcb.14792
Shipley B (2009) Confirmatory path analysis in a generalized multilevel context. Ecology 90:363–368. https://doi.org/10.1890/08-1034.1
Slik JWF, Paoli G, Mcguire K et al (2013) Large trees drive forest aboveground biomass variation in moist lowland forests across the tropics. Glob Ecol Biogeogr 22:1261–1271. https://doi.org/10.1111/geb.12092
Tilman D (2001) Functional diversity. Encycl Biodivers 3:109–120. https://doi.org/10.1016/B0-12-226865-2/00132-2
Tilman D, Lehman CL, Thomson KT (1997) Plant diversity and ecosystem productivity: theoretical considerations. Proc Natl Acad Sci 94:1857–1861. https://doi.org/10.1073/pnas.94.5.1857
Van Der Sande MT, Peña-Claros M, Ascarrunz N et al (2017) Abiotic and biotic drivers of biomass change in a Neotropical forest. J Ecol 105:1223–1234. https://doi.org/10.1111/1365-2745.12756
Wang S, Jiménez-Alfaro B, Pan S et al (2021) Differential responses of forest strata species richness to paleoclimate and forest structure. For Ecol Manage 499:119605. https://doi.org/10.1016/j.foreco.2021.119605
Whittaker RH (1972) Evolution and measurement of species diversity. Taxon 21:213–251. https://doi.org/10.2307/1218190
Wright JS (2002) Plant diversity in tropical forests: a review of mechanisms of species coexistence. Oecologia 130:1–14. https://doi.org/10.1007/s004420100809
Xu G, Chen H, Shi Z et al (2021) Mycorrhizal and rhizospheric fungal community assembly differs during subalpine forest restoration on the eastern Qinghai-Tibetan Plateau. Plant Soil 458:245–259. https://doi.org/10.1007/s11104-019-04400-7
Yachi S, Loreau M (2007) Does complementary resource use enhance ecosystem functioning? A model of light competition in plant communities. Ecol Lett 10:54–62. https://doi.org/10.1111/j.1461-0248.2006.00994.x
Yu K, Chen X, Pan G et al (2016) Dynamics of soil available phosphorus and its impact factors under simulated climate change in typical farmland of Taihu Lake region, China. Environ Monit Assess 188:1–8. https://doi.org/10.1007/s10661-015-5087-0
Yuan Z, Wang S, Gazol A et al (2016) Multiple metrics of diversity have different effects on temperate forest functioning over succession. Oecologia 182:1175–1185. https://doi.org/10.1007/s00442-016-3737-8
Yuan Z, Wang S, Ali A et al (2018) Aboveground carbon storage is driven by functional trait composition and stand structural attributes rather than biodiversity in temperate mixed forests recovering from disturbances. Ann for Sci 75:1–13. https://doi.org/10.1007/s13595-018-0745-3
Zemunik G, Turner BL, Lambers H et al (2016) Increasing plant species diversity and extreme species turnover accompany declining soil fertility along a long-term chronosequence in a biodiversity hotspot. J Ecol 104:792–805. https://doi.org/10.1111/1365-2745.12546
Zhang Y, Chen HYH (2015) Individual size inequality links forest diversity and above-ground biomass. J Ecol 103:1245–1252. https://doi.org/10.1111/1365-2745.12425
Zhang Y, Chen HYH, Taylor AR (2016) Aboveground biomass of understorey vegetation has a negligible or negative association with overstorey tree species diversity in natural forests. Glob Ecol Biogeogr 25:141–150. https://doi.org/10.1111/geb.12392
Zhang Y, Chen HYH, Taylor AR (2017) Positive species diversity and above-ground biomass relationships are ubiquitous across forest strata despite interference from overstorey trees. Funct Ecol 31:419–426. https://doi.org/10.1111/1365-2435.12699
Zheng Q, Hu Y, Zhang S et al (2019) Soil multifunctionality is affected by the soil environment and by microbial community composition and diversity. Soil Biol Biochem 136:107521. https://doi.org/10.1016/j.soilbio.2019.107521
Acknowledgements
The work was supported by the Fundamental Research Funds of CAF (CAFYBB2021ZA002-2, CAFYBB2022QC002, CAFYBB2022SY021, and CAFYBB2022SY024), the National Key Research and Development Program of China (2021YFD2200405) and the National Natural Science Foundation of China (32201321).
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ZS conceived the study and designed the methodology; FL, SL, MC, GX, JC, and HX collected the data; and FL analyzed the data and wrote the manuscript. ZS, GX, and SL reviewed the manuscript. All authors contributed to drafts and agree final approval for the published version of the manuscript.
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Li, F., Shi, Z., Liu, S. et al. Stand structural diversity and edaphic properties regulate aboveground biomass of Abies fargesii var. faxoniana primary forest on the eastern Qinghai-Tibetan Plateau. Eur J Forest Res (2024). https://doi.org/10.1007/s10342-024-01697-7
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DOI: https://doi.org/10.1007/s10342-024-01697-7