{
  "abstract": "Forests are a major terrestrial carbon sink, and higher overstory diversity is often associated with greater carbon accrual in trees and soils. Yet the contribution of the understory vegetation, the most species-rich forest stratum, to tree and soil carbon accumulation remains largely unknown. Here, we analyzed Canada’s National Forest Inventory database to explore whether understory diversity (richness and evenness of shrubs, herbs, and bryophytes) is associated with carbon accumulation in overstory trees and soils across environmental gradients of air temperature, climate moisture, and nitrogen deposition. We found that herb and bryophyte species richness were positively associated with decadal tree carbon accumulation, with the strongest effects at warmer, drier, and nitrogen-limited sites. In contrast, understory diversity showed no consistent association with soil carbon accumulation. On a decadal scale, increasing herb and bryophyte species richness from their minimum to median values corresponded to 15% and 6% increases in tree carbon accumulation, respectively. These results provide large-scale observational evidence that diversity in lower vegetation layers is positively associated with overstory carbon sequestration and reveal strong environmental modulation of these multistratal biodiversity–ecosystem functioning relationships. These findings suggest that species-rich herb and bryophyte communities should be considered in strategies for sustaining forest carbon sinks, especially under warmer, drier conditions and nutrient limitation.",
  "abstract_shown": true,
  "affiliations": [
    "State Key Laboratory for Development and Utilization of Forest Food Resources, Zhejiang Agriculture and Forestry University, Hangzhou 311300, China",
    "Key Laboratory of Carbon Sequestration and Emission Reduction in Agriculture and Forestry of Zhejiang Province, College of Environmental and Resources Sciences, Zhejiang Agriculture and Forestry University, Hangzhou 311300, China",
    "Department of Renewable Resources, University of Alberta, Edmonton, AB T6G 2E3, Canada",
    "College of Grassland Science, Inner Mongolia Agricultural University, Hohhot 010011, China",
    "Institute for Global Change Biology, and School for Environment and Sustainability, University of Michigan, Ann Arbor, MI 48109",
    "Faculty of Natural Resources Management, Lakehead University, Thunder Bay, ON P7B 5E1, Canada",
    "Faculty of Forestry and Environmental Management, University of New Brunswick, Fredericton, NB E3B 5A3, Canada",
    "Graduate School of Advanced Science and Engineering, Hiroshima University, Hiroshima 739-8529, Japan",
    "Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China",
    "Appalachian Laboratory, University of Maryland Center for Environmental Science, Frostburg, MD 21532",
    "Department of Forest Resources, University of Minnesota, St. Paul, MN 55108",
    "Hawkesbury Institute for the Environment, Western Sydney University, Penrith, NSW 2753, Australia"
  ],
  "article_type": "research article",
  "arxiv": null,
  "author_total": 13,
  "authors": [
    {
      "affs": [
        0,
        1
      ],
      "name": "Yakun Zhang",
      "orcid": "0000-0002-2917-2637"
    },
    {
      "affs": [
        0
      ],
      "name": "Rui Zhang",
      "orcid": "0000-0002-7235-505X"
    },
    {
      "affs": [
        2
      ],
      "name": "Scott X. Chang",
      "orcid": "0000-0002-7624-439X"
    },
    {
      "affs": [
        3,
        4,
        5
      ],
      "name": "Han Y. H. Chen",
      "orcid": "0000-0001-9477-5541"
    },
    {
      "affs": [
        6
      ],
      "name": "Anthony R. Taylor",
      "orcid": "0000-0002-2122-6792"
    },
    {
      "affs": [
        7
      ],
      "name": "Masumi Hisano",
      "orcid": "0000-0002-3869-8542"
    },
    {
      "affs": [
        8
      ],
      "name": "Chen Chen",
      "orcid": "0000-0002-8340-6115"
    },
    {
      "affs": [
        8
      ],
      "name": "Sai Peng",
      "orcid": "0000-0002-0237-0684"
    },
    {
      "affs": [
        0,
        1
      ],
      "name": "Ya Wang",
      "orcid": "0009-0001-1176-2583"
    },
    {
      "affs": [
        9
      ],
      "name": "Xin Chen",
      "orcid": "0000-0003-0801-8566"
    },
    {
      "affs": [
        0
      ],
      "name": "Jiasheng Wu",
      "orcid": "0000-0002-2515-4677"
    },
    {
      "affs": [
        0,
        1,
        2,
        3
      ],
      "name": "Xinli Chen",
      "orcid": "0000-0003-0542-5959"
    },
    {
      "affs": [
        4,
        10,
        11
      ],
      "name": "Peter B. Reich",
      "orcid": "0000-0003-4424-662X"
    }
  ],
  "bibtex": "https://research.draconitic.com/paper/8dfan0jw8etg0.bib",
  "cited_by": 0,
  "doi": "10.1073/pnas.2615624123",
  "fulltext_url": "https://pmc.ncbi.nlm.nih.gov/articles/PMC13715778/",
  "id": "8dfan0jw8etg0",
  "issns": [
    "0027-8424",
    "1091-6490"
  ],
  "landing_url": "https://doi.org/10.1073/pnas.2615624123",
  "language": "en",
  "licence": "CC-BY-NC-ND-4.0",
  "licence_label": "CC BY-NC-ND 4.0",
  "licence_source": "article",
  "licence_url": "https://creativecommons.org/licenses/by-nc-nd/4.0/",
  "metadata_licence": null,
  "native_subjects": [
    [
      "heading",
      "414"
    ],
    [
      "heading",
      "Biological Sciences"
    ],
    [
      "heading",
      "Ecology"
    ],
    [
      "heading",
      "Sdg-15-Life-on-Land"
    ],
    [
      "openalex",
      "2309|Forest ecology and management"
    ]
  ],
  "openalex": "W7222917974",
  "pdf_url": null,
  "pmcid": "PMC13715778",
  "pmid": "42896055",
  "published": "2026-10-19",
  "publisher": "National Academy of Sciences",
  "ris": "https://research.draconitic.com/paper/8dfan0jw8etg0.ris",
  "sources": [
    {
      "first_seen": "2026-11-01T12:22:59Z",
      "native_id": "PMC13715778",
      "source": "pmc",
      "url": "https://doi.org/10.1073/pnas.2615624123"
    }
  ],
  "subjects": [
    {
      "field": "biology",
      "origin": "heading:Ecology",
      "subfield": "ecology"
    }
  ],
  "title": "Understory diversity is positively associated with overstory carbon sequestration",
  "title_text": "Understory diversity is positively associated with overstory carbon sequestration",
  "type": "peer-reviewed",
  "updated_at": "2026-11-01T12:22:59Z",
  "url": "https://research.draconitic.com/paper/8dfan0jw8etg0",
  "venue": "Proceedings of the National Academy of Sciences of the United States of America",
  "versions": []
}