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Peer-reviewedCC BY-NC-ND 4.0Proceedings of the National Academy of Sciences of the United States of America

Understory diversity is positively associated with overstory carbon sequestration

  • Yakun Zhang1,2
  • Rui Zhang1
  • Scott X. Chang3
  • Han Y. H. Chen4,5,6
  • Anthony R. Taylor7
  • Masumi Hisano8
  • Chen Chen9
  • Sai Peng9
  • Ya Wang1,2
  • Xin Chen10
  • Jiasheng Wu1
  • Xinli Chen1,2,3,4
1 more authors
  • Peter B. Reich5,11,12
12 affiliations
  1. 1State Key Laboratory for Development and Utilization of Forest Food Resources, Zhejiang Agriculture and Forestry University, Hangzhou 311300, China
  2. 2Key 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
  3. 3Department of Renewable Resources, University of Alberta, Edmonton, AB T6G 2E3, Canada
  4. 4College of Grassland Science, Inner Mongolia Agricultural University, Hohhot 010011, China
  5. 5Institute for Global Change Biology, and School for Environment and Sustainability, University of Michigan, Ann Arbor, MI 48109
  6. 6Faculty of Natural Resources Management, Lakehead University, Thunder Bay, ON P7B 5E1, Canada
  7. 7Faculty of Forestry and Environmental Management, University of New Brunswick, Fredericton, NB E3B 5A3, Canada
  8. 8Graduate School of Advanced Science and Engineering, Hiroshima University, Hiroshima 739-8529, Japan
  9. 9Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
  10. 10Appalachian Laboratory, University of Maryland Center for Environmental Science, Frostburg, MD 21532
  11. 11Department of Forest Resources, University of Minnesota, St. Paul, MN 55108
  12. 12Hawkesbury Institute for the Environment, Western Sydney University, Penrith, NSW 2753, Australia
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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.

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Cite

BibTeXRIS
BibTeX
@article{zhang2026understory,
  title = {{Understory diversity is positively associated with overstory carbon sequestration}},
  author = {Yakun Zhang and Rui Zhang and Scott X. Chang and Han Y. H. Chen and Anthony R. Taylor and Masumi Hisano and Chen Chen and Sai Peng and Ya Wang and Xin Chen and Jiasheng Wu and Xinli Chen and Peter B. Reich},
  journal = {Proceedings of the National Academy of Sciences of the United States of America},
  year = {2026},
  month = oct,
  doi = {10.1073/pnas.2615624123},
  url = {https://doi.org/10.1073/pnas.2615624123},
  note = {CC BY-NC-ND 4.0},
}
RIS
TY  - JOUR
TI  - Understory diversity is positively associated with overstory carbon sequestration
AU  - Yakun Zhang
AU  - Rui Zhang
AU  - Scott X. Chang
AU  - Han Y. H. Chen
AU  - Anthony R. Taylor
AU  - Masumi Hisano
AU  - Chen Chen
AU  - Sai Peng
AU  - Ya Wang
AU  - Xin Chen
AU  - Jiasheng Wu
AU  - Xinli Chen
AU  - Peter B. Reich
PY  - 2026
DA  - 2026/10/19
JO  - Proceedings of the National Academy of Sciences of the United States of America
DO  - 10.1073/pnas.2615624123
UR  - https://doi.org/10.1073/pnas.2615624123
AB  - 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.
SN  - 0027-8424
SN  - 1091-6490
ER  -