{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T07:01:36Z","timestamp":1790146896598,"version":"4.0.1"},"reference-count":61,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:p>Forests in karst landscapes occur across strong edaphic gradients that may alter biodiversity\u2013carbon relationships, yet these relationships are commonly evaluated using tree aboveground biomass carbon, potentially obscuring contrasting responses among forest strata and biomass compartments. We examined how soil fertility context modifies multidimensional biodiversity\u2013carbon associations across aboveground and allometrically estimated belowground compartments in a subtropical karst forest landscape of China. Fifty-four plots were classified as low-fertility (LF) or high-fertility (HF) based on soil nutrient conditions. We quantified tree and shrub aboveground carbon and allometrically estimated root carbon, and assessed plot-level taxonomic diversity, functional diversity, community-weighted trait composition, structural diversity, and abiotic conditions. Associations were evaluated using correlation, regression, relative-importance, and piecewise structural equation analyses. Tree aboveground and estimated tree root carbon were higher under LF conditions, whereas both shrub carbon components were higher under HF conditions. Associations between plot-level biodiversity attributes and carbon stocks varied strongly among woody strata and fertility contexts. Under HF conditions, plot-level species diversity was positively associated with tree carbon but negatively associated with shrub carbon, while structural diversity was positively related to estimated tree root carbon but negatively related to shrub carbon. Under LF conditions, plot-level functional diversity showed negative fitted paths to tree carbon but positive paths to shrub carbon. Predictor rankings and pathway configurations differed across all four carbon compartments, and the measured predictors explained more variation under HF than LF conditions. These findings show that soil fertility is associated with contrasting carbon-storage patterns between tree and shrub strata and that aggregated metrics can conceal reversals in biodiversity\u2013carbon relationships. Resolving forest strata and aboveground and allometrically estimated root-carbon components is essential for understanding plant diversity and carbon storage and informing vegetation restoration in karst landscapes.<\/jats:p>","DOI":"10.3389\/fpls.2026.1942459","type":"journal-article","created":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T06:18:33Z","timestamp":1790144313000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Soil fertility context shapes biodiversity\u2013carbon associations across forest strata in a subtropical karst landscape"],"prefix":"10.3389","volume":"17","author":[{"given":"Jie","family":"Zheng","sequence":"first","affiliation":[{"name":"Yunnan Collaborative Innovation Center for Plateau Lake Ecology and Environmental Health, College of Agronomy and Life Sciences, Kunming University, Kunming","place":["Yunnan, China"]},{"name":"Key Laboratory of Eco-Environments in the Three Gorges Reservoir Region (Ministry of Education), College of Life Sciences, Southwest University","place":["Chongqing, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jufu","family":"Zhang","sequence":"additional","affiliation":[{"name":"Yunnan Collaborative Innovation Center for Plateau Lake Ecology and Environmental Health, College of Agronomy and Life Sciences, Kunming University, Kunming","place":["Yunnan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Meigui","family":"Yang","sequence":"additional","affiliation":[{"name":"Yunnan Collaborative Innovation Center for Plateau Lake Ecology and Environmental Health, College of Agronomy and Life Sciences, Kunming University, Kunming","place":["Yunnan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ailian","family":"Zou","sequence":"additional","affiliation":[{"name":"Yunnan Collaborative Innovation Center for Plateau Lake Ecology and Environmental Health, College of Agronomy and Life Sciences, Kunming University, Kunming","place":["Yunnan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hao","family":"Xiong","sequence":"additional","affiliation":[{"name":"Yunnan Collaborative Innovation Center for Plateau Lake Ecology and Environmental Health, College of Agronomy and Life Sciences, Kunming University, Kunming","place":["Yunnan, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Muhammad","family":"Arif","sequence":"additional","affiliation":[{"name":"School of Economics and Business Administration, Heilongjiang University","place":["Harbin, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,23]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.foreco.2019.01.007","article-title":"Multiple abiotic and biotic drivers of aboveground biomass shift with forest stratum","volume":"436","author":"Ali","year":"2019","journal-title":"For. Ecol. Manage."},{"key":"B2","doi-asserted-by":"publisher","first-page":"1097","DOI":"10.1038\/s41467-022-28748-0","article-title":"Tree functional traits, forest biomass, and tree species diversity interact with site properties to drive forest soil carbon","volume":"13","author":"Augusto","year":"2022","journal-title":"Nat. Commun."},{"key":"B3","doi-asserted-by":"publisher","first-page":"1976","DOI":"10.1038\/s41467-026-68668-x","article-title":"European forest carbon and biodiversity policies have a limited win-win potential","volume":"17","author":"Balducci","year":"2026","journal-title":"Nat. Commun."},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.1002\/ecs2.2185","article-title":"Forests, shrubs, and terrain: top-down and bottom-up controls on forest structure","volume":"9","author":"Bolstad","year":"2018","journal-title":"Ecosphere"},{"key":"B5","doi-asserted-by":"publisher","first-page":"112","DOI":"10.1016\/j.pecon.2023.02.003","article-title":"No relationship between biodiversity and forest carbon sink across the subtropical Brazilian Atlantic Forest","volume":"21","author":"Bordin","year":"2023","journal-title":"Perspect. Ecol. Conserv."},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0151852","article-title":"Tree size inequality reduces forest productivity: an analysis combining inventory data for ten European species and a light competition model","volume":"11","author":"Bourdier","year":"2016","journal-title":"PloS One"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1093\/nsr\/nwaf093","article-title":"Understory shrub diversity: equally vital as overstory tree diversity to promote forest productivity","volume":"12","author":"Chen","year":"2025","journal-title":"Natl. Sci. Rev."},{"key":"B8","doi-asserted-by":"publisher","first-page":"8615","DOI":"10.1038\/s41467-024-53004-y","article-title":"Resource availability enhances positive tree functional diversity effects on carbon and nitrogen accrual in natural forests","volume":"15","author":"Chen","year":"2024","journal-title":"Nat. Commun."},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.48130\/FR-2023-0006","article-title":"Forest understory vegetation study: current status and future trends","volume":"3","author":"Deng","year":"2023","journal-title":"For. Res."},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1016\/j.tfp.2025.100973","article-title":"Mass-ratio effects outperform niche complementarity in predicting aboveground carbon storage in southeastern United States pine and mixed forests","volume":"21","author":"Fatunsin","year":"2025","journal-title":"Trees For. People"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1016\/j.foreco.2025.122719","article-title":"Structural diversity enhances carbon storage in mixed oak-pine forests of the Southeast United States","volume":"586","author":"Fatunsin","year":"2025","journal-title":"For. Ecol. Manage."},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1016\/j.catena.2025.109069","article-title":"Decoding soil carbon and nitrogen dynamics: the integrated role of biotic and abiotic drivers in subtropical forests","volume":"256","author":"Gao","year":"2025","journal-title":"CATENA"},{"key":"B13","doi-asserted-by":"publisher","first-page":"16806","DOI":"10.1002\/ece3.8312","article-title":"Mass-ratio and complementarity effects simultaneously drive aboveground biomass in temperate Quercus forests through stand structure","volume":"11","author":"Gao","year":"2021","journal-title":"Ecol. Evol."},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.1016\/j.ecolind.2024.112634","article-title":"Characteristics of forest understory herbaceous vegetation and its influencing factors in biodiversity hotspots in China","volume":"167","author":"Geng","year":"2024","journal-title":"Ecol. Indic."},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1093\/jpe\/rtaf100","article-title":"Structural diversity drives forest biomass along altitudinal gradients in subtropical forests","volume":"18","author":"Guo","year":"2025","journal-title":"J. Plant Ecol."},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1016\/j.ecolind.2024.111685","article-title":"Biodiversity and carbon stocks of the understory vegetation as indicators for forest health in the Zabarwan Mountain Range, Indian Western Himalaya","volume":"159","author":"Haq","year":"2024","journal-title":"Ecol. Indic."},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1111\/1365-2745.70200","article-title":"Structural diversity shifts from negative to positive associations with forest productivity via basal area, stand age, and precipitation thresholds","volume":"114","author":"Hwang","year":"2026","journal-title":"J. Ecol."},{"key":"B18","doi-asserted-by":"publisher","first-page":"1756","DOI":"10.1111\/1365-2435.13825","article-title":"Above- and below-ground complementarity rather than selection drive tree diversity\u2013productivity relationships in European forests","volume":"35","author":"Jing","year":"2021","journal-title":"Funct. Ecol."},{"key":"B19","doi-asserted-by":"publisher","first-page":"653","DOI":"10.1016\/j.pld.2025.04.008","article-title":"Tree-based attributes of large trees more effectively regulate aboveground carbon stock than trait-based ones in temperate deciduous forests","volume":"47","author":"Kazempour Larsary","year":"2025","journal-title":"Plant Divers."},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1016\/j.ecolind.2025.113131","article-title":"How do leaf functional traits influence above-ground tree carbon in tropical hill forests of Bangladesh","volume":"171","author":"Khan","year":"2025","journal-title":"Ecol. Indic."},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1016\/j.fecs.2025.100386","article-title":"Variation of above-ground tree biomass and soil carbon stocks across neotropical forest types","volume":"15","author":"Lanuza","year":"2026","journal-title":"For. Ecosyst."},{"key":"B22","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1111\/2041-210X.12512","article-title":"piecewiseSEM: Piecewise structural equation modelling in r for ecology, evolution, and systematics","volume":"7","author":"Lefcheck","year":"2016","journal-title":"Methods Ecol. Evol."},{"key":"B23","doi-asserted-by":"publisher","first-page":"684","DOI":"10.1016\/j.scitotenv.2018.11.024","article-title":"Drivers of tree carbon storage in subtropical forests","volume":"654","author":"Li","year":"2019","journal-title":"Sci. Total Environ."},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1016\/j.foreco.2025.123309","article-title":"Context-dependent forest carbon storage: a shift in dominance from structural complexity to large-sized trees across disturbance gradients","volume":"600","author":"Li","year":"2026","journal-title":"For. Ecol. Manage."},{"key":"B25","doi-asserted-by":"publisher","first-page":"399","DOI":"10.1016\/j.pld.2025.12.005","article-title":"Tree structural diversity mediates vegetation carbon storage in dry-hot valley savannas along an elevational gradient","volume":"48","author":"Li","year":"2026","journal-title":"Plant Divers."},{"key":"B26","doi-asserted-by":"publisher","first-page":"435","DOI":"10.1007\/s11104-016-2976-0","article-title":"Traits of dominant tree species predict local scale variation in forest aboveground and topsoil carbon stocks","volume":"409","author":"Lin","year":"2016","journal-title":"Plant Soil"},{"key":"B27","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1186\/s13021-026-00437-1","article-title":"Research progress on the interaction between forest carbon sequestration and climate change","volume":"21","author":"Lin","year":"2026","journal-title":"Carbon Balance Manage."},{"key":"B28","doi-asserted-by":"publisher","first-page":"1688","DOI":"10.1111\/1365-2435.14334","article-title":"Environmental filtering drives biodiversity\u2013spatial stability relationships in a large temperate forest region","volume":"37","author":"Lin","year":"2023","journal-title":"Funct. Ecol."},{"key":"B29","doi-asserted-by":"publisher","first-page":"2786","DOI":"10.1002\/ldr.5531","article-title":"Exploring the spatial relationship between carbon storage and biodiversity: a systematic review and meta-analysis","volume":"36","author":"Liu","year":"2025","journal-title":"Land Degrad. Dev."},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.1002\/eap.70211","article-title":"Structural characteristics mediate forest mitigation potential against climate change and biodiversity loss","volume":"36","author":"Lunow","year":"2026","journal-title":"Ecol. Appl."},{"key":"B31","doi-asserted-by":"publisher","first-page":"2174","DOI":"10.1111\/jbi.13669","article-title":"Forest community assembly is driven by different strata-dependent mechanisms along an elevational gradient","volume":"46","author":"Luo","year":"2019","journal-title":"J. Biogeogr."},{"key":"B32","doi-asserted-by":"publisher","first-page":"1449","DOI":"10.1111\/ele.13330","article-title":"Greater than the sum of the parts: how the species composition in different forest strata influence ecosystem function","volume":"22","author":"Luo","year":"2019","journal-title":"Ecol. Lett."},{"key":"B33","doi-asserted-by":"publisher","first-page":"1037","DOI":"10.1111\/1365-2745.14076","article-title":"The interactive effects of soil fertility and tree mycorrhizal association explain spatial variation of diversity\u2013biomass relationships in a subtropical forest","volume":"111","author":"Ma","year":"2023","journal-title":"J. Ecol."},{"key":"B34","doi-asserted-by":"publisher","first-page":"475","DOI":"10.1111\/nph.15742","article-title":"Tree mycorrhizal associations mediate soil fertility effects on forest community structure in a temperate forest","volume":"223","author":"Mao","year":"2019","journal-title":"New Phytol."},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1002\/ecm.1568","article-title":"Scale-dependent diversity\u2013biomass relationships can be driven by tree mycorrhizal association and soil fertility","volume":"93","author":"Mao","year":"2023","journal-title":"Ecol. Monogr."},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1016\/j.jenvman.2023.118929","article-title":"Biodiversity and carbon conservation under the ecosystem stability of tropical forests","volume":"345","author":"Maure","year":"2023","journal-title":"J. Environ. Manage."},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1016\/j.ecolind.2025.113990","article-title":"Interactive effects of functional, taxonomic, and structural diversity on aboveground carbon storage in humid subtropical mixed forests","volume":"178","author":"Naka","year":"2025","journal-title":"Ecol. Indic."},{"key":"B38","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1111\/geb.13800","article-title":"Both the selection and complementarity effects underpin the effect of structural diversity on aboveground biomass in tropical forests","volume":"33","author":"Noul\u00e8koun","year":"2024","journal-title":"Global Ecol. Biogeogr."},{"key":"B39","volume-title":"R: A Language and Environment for Statistical Computing","year":"2023"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.70089","article-title":"Tree diversity increases carbon stocks and fluxes above\u2014but not belowground in a tropical forest experiment","volume":"31","author":"Schnabel","year":"2025","journal-title":"Global Change Biol."},{"key":"B41","doi-asserted-by":"publisher","first-page":"5321","DOI":"10.1111\/gcb.16697","article-title":"Carbon\u2013biodiversity relationships in a highly diverse subtropical forest","volume":"29","author":"Schuldt","year":"2023","journal-title":"Global Change Biol."},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1016\/j.tfp.2026.101244","article-title":"The stand structure, species diversity, and carbon of Mendawak Protected Forest, West Kalimantan-Indonesia","volume":"25","author":"Siregar","year":"2026","journal-title":"Trees For. People"},{"key":"B43","doi-asserted-by":"publisher","first-page":"1848","DOI":"10.1038\/s41598-018-37190-6","article-title":"Both mass ratio effects and community diversity drive biomass production in a grassland experiment","volume":"9","author":"Sonkoly","year":"2019","journal-title":"Sci. Rep."},{"key":"B44","doi-asserted-by":"publisher","first-page":"2302","DOI":"10.1038\/s41598-020-58913-8","article-title":"Soil properties explain tree growth and mortality, but not biomass, across phosphorus-depleted tropical forests","volume":"10","author":"Soong","year":"2020","journal-title":"Sci. Rep."},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1016\/j.foreco.2024.122328","article-title":"Identifying the stand properties that support both high biodiversity and carbon storage in German forests","volume":"572","author":"Springer","year":"2024","journal-title":"For. Ecol. Manage."},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1016\/j.ecolind.2025.114398","article-title":"Forest structural attributes as indicators and mediators of aboveground carbon storage and woody plant species richness in natural forests","volume":"181","author":"Sun","year":"2025","journal-title":"Ecol. Indic."},{"key":"B47","doi-asserted-by":"publisher","first-page":"706","DOI":"10.1016\/j.ecofro.2025.11.002","article-title":"Assessing tree species diversity, aboveground biomass, and soil fertility across forest types of the eastern Hindukush Mountains","volume":"46","author":"Ullah","year":"2026","journal-title":"Ecol. Front."},{"key":"B48","doi-asserted-by":"publisher","first-page":"461","DOI":"10.1111\/1365-2435.12968","article-title":"Soil fertility and species traits, but not diversity, drive productivity and biomass stocks in a Guyanese tropical rainforest","volume":"32","author":"van der Sande","year":"2018","journal-title":"Funct. Ecol."},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.1016\/j.pld.2026.05.011","article-title":"Resource-poor environments drive more prevalent habitat associations in Ectomycorrhizal tree species: Evidence from tropical, subtropical, and subalpine forest dynamics plots","author":"Wan","year":"2026","journal-title":"Plant Divers"},{"key":"B50","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1111\/nph.17590","article-title":"An integrated framework of plant form and function: the belowground perspective","volume":"232","author":"Weigelt","year":"2021","journal-title":"New Phytol."},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.1016\/j.scitotenv.2022.155907","article-title":"Tropical forest strata shifts in plant structural diversity-aboveground carbon relationships along altitudinal gradients","volume":"838","author":"Wen","year":"2022","journal-title":"Sci. Total Environ."},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.70120","article-title":"Functional diversity explains ecosystem carbon storage in subtropical forests","volume":"31","author":"Wu","year":"2025","journal-title":"Global Change Biol."},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1002\/ecy.4269","article-title":"Structural diversity is better associated with forest productivity than species or functional diversity","volume":"105","author":"Zhai","year":"2024","journal-title":"Ecology"},{"key":"B54","doi-asserted-by":"publisher","first-page":"419","DOI":"10.1111\/1365-2435.12699","article-title":"Positive species diversity and above-ground biomass relationships are ubiquitous across forest strata despite interference from overstorey trees","volume":"31","author":"Zhang","year":"2017","journal-title":"Funct. Ecol."},{"key":"B55","doi-asserted-by":"publisher","first-page":"1206","DOI":"10.1111\/gcb.16540","article-title":"Soil fertility shifts the relative importance of saprotrophic and mycorrhizal fungi for maintaining ecosystem stability","volume":"29","author":"Zhang","year":"2023","journal-title":"Global Change Biol."},{"key":"B56","doi-asserted-by":"publisher","DOI":"10.1016\/j.scitotenv.2022.160548","article-title":"Drivers of aboveground biomass shift with forest stratum in temperate forest of North China","volume":"860","author":"Zhang","year":"2023","journal-title":"Sci. Total Environ."},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.1093\/jpe\/rtae043","article-title":"Multi-dimensionality in plant root traits: progress and challenges","volume":"17","author":"Zhao","year":"2024","journal-title":"J. Plant Ecol."},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1016\/j.pld.2026.05.003","article-title":"Latitudinal diversity gradients of soil seed bank and aboveground vegetation in China","author":"Zhao","year":"2026","journal-title":"Plant Divers"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1016\/j.envres.2025.121101","article-title":"Flooding drives plant diversity\u2013biomass relationships in riparian zones of the three Gorges reservoir area","volume":"271","author":"Zheng","year":"2025","journal-title":"Environ. Res."},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2025.1728262","article-title":"Synergies and trade-offs in forest carbon pools: separating universal drivers from forest type-specific controls","volume":"16","author":"Zhou","year":"2026","journal-title":"Front. Plant Sci."},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1016\/j.fecs.2024.100187","article-title":"No generality in biodiversity-productivity relationships along elevation in temperate and subtropical forest landscapes","volume":"11","author":"Zou","year":"2024","journal-title":"For. Ecosyst."}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1942459\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T06:18:34Z","timestamp":1790144314000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1942459\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,23]]},"references-count":61,"alternative-id":["10.3389\/fpls.2026.1942459"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2026.1942459","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,23]]},"article-number":"1942459"}}