Abstract
Large, old trees store a considerable part of the total carbon in old-growth forests; however, their influence on long-term climate change mitigation capacity remains unclear. Damage caused by wind and insect outbreaks can rapidly kill trees, and these natural events are expected to increase with climate change, thereby reducing the long-term carbon storage capacity of old-growth stands. The aim was to estimate the importance of large, old trees in carbon storage at the stand level and the potential risks for the carbon stock if they die. The data were collected from Scots pine and Norway spruce stands on mineral soils, of which 44 were old-growth stands, and 47 were mature stands two times younger than the former. Stand-level carbon storage in tree biomass was investigated; the presence of deadwood was analyzed. The carbon stock changes were calculated by excluding the 1–15 largest trees per sample plot. The results showed that the largest trees in old-growth coniferous stands hold a disproportionally large (~ 50% of the total tree biomass carbon pool) share of carbon compared to mature stands, ~ 50% and ~ 10%, respectively. A reduction of the tree biomass carbon pool by approximately 50% required a removal of around 30% more trees in mature than in old-growth stands. A relatively low proportion of deadwood from the total stand volume (~ 20%) suggests a limited single-tree mortality and vulnerable mitigation potential in the assessed old-growth stands. Nevertheless, continuous single-tree mortality due to ageing or windthrow re-shapes forest structure, and raises the potential risk of the stands to maintain and/or further increase substantial tree biomass-linked carbon pools.
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Data availability
The datasets generated during and/or analyzed during the current study are available from the corresponding author.
References
Berglund, H. & Kuuluvainen, T. Representative boreal forest habitats in Northern Europe, and a revised model for ecosystem management and biodiversity conservation. Ambio 50, 1003–1017. https://doi.org/10.1007/s13280-020-01444-3 (2021).
Kilpeläinen, A. & Peltola, H. Carbon sequestration and storage in European forests. In Forest Bioeconomy and Climate Change. Managing Forest Ecosystems Vol. 42 (eds Hetemäki, L. et al.) (Springer, 2022). https://doi.org/10.1007/978-3-030-99206-4_6.
Barredo, J. et al. Mapping and Assessment of Primary and old-growth Forests in Europe (EUR 30661 EN, 2021). https://doi.org/10.2760/797591
Matuszkiewicz, J. M., Affek, A. N. & Kowalska, A. Current and potential carbon stock in the forest communities of the Białowieża biosphere reserve. Ecol. Manag. 502, 119702. https://doi.org/10.1016/j.foreco.2021.119702 (2021).
Sabatini, F. M. et al. European primary forest database v2.0. Sci. Data. 8, 220. https://doi.org/10.1038/s41597-021-00988-7 (2021).
Motta, R. et al. Old-growth forests in the dinaric alps of Bosnia-Herzegovina and montenegro: a continental hot-spot for research and biodiversity. Front. Glob Change. 7, 01–16. https://doi.org/10.3389/ffgc.2024.1371144 (2024).
Buchwald, E. A hierarchical terminology for more or less natural forests in relation to sustainable management and biodiversity conservation. Third Expert Meeting on Harmonizing Forest-related Definitions. FAO. Proceedings at (2005). https://forestsandco.files.wordpress.com/2015/11/buchwald_2002_definitions.pdf
Martin, M., Shorohova, E. & Fenton, N. J. Embracing the complexity and the richness of boreal old-growth forests: A further step toward their ecosystem management. In Boreal Forests in the Face of Climate Change Vol. 74 (eds Girona, M. M. et al.) (Springer, 2023). https://doi.org/10.1007/978-3-031-15988-6_7.
O’Brien, L. et al. Protecting old-growth Forests in Europe – a Review of Scientific Evidence To Inform Policy Implementation (Final report of European Forest Institute at, 2021). https://doi.org/10.36333/rs1
Barredo, J. I., Vizzarri, M. & Kužnelová, K. Archetypal typology of European forest ecosystems integrating management intensity and naturalness. Ambio 53, 1587–1598. https://doi.org/10.1007/s13280-024-02050-3 (2024).
Pardos, M. et al. Carbon storage potentiality in successional and secondary old growth forests. Eur. J. Res. 144, 193–211. https://doi.org/10.1007/s10342-024-01750-5 (2025).
Taylor, A. R., Seedre, M., Brassard, B. W. & Chen, H. Y. Decline in net ecosystem productivity following canopy transition to late-succession forests. Ecosystems 17, 778–791. https://doi.org/10.1007/s10021-014-9759-3 (2014).
Cerioni, M. et al. Recovery and resilience of European temperate forests after large and severe disturbances. Glob Change Biol. 30 (2), e17159. https://doi.org/10.1111/gcb.17159 (2023).
Kuuluvainen, T. & Gauthier, S. Young and old forest in the boreal: critical stages of ecosystem dynamics and management under global change. Ecosyst. 5, 26. https://doi.org/10.1186/s40663-018-0142-2 (2018).
Seidl, R. et al. Globally consistent climate sensitivity of natural disturbances across boreal and temperate forest ecosystems. Ecography 43 (7), 967–978. https://doi.org/10.1111/ecog.04995 (2020).
Martin-Benito, D., Molina-Valero, J. A., Pérez-Cruzado, C., Bigler, C. & Bugmann, H. Development and long-term dynamics of old-growth beech-fir forests in the pyrenees: evidence from dendroecology and dynamic vegetation modelling. Ecol. Manag. 524, 120541. https://doi.org/10.1016/j.foreco.2022.120541 (2022).
Keith, H. et al. Carbon carrying capacity in primary forests shows potential for mitigation achieving the European green deal 2030 target. Commun. Earth Environ. 5, 256. https://doi.org/10.1038/s43247-024-01416-5 (2024).
Anderson, S., Knapp, B. O. & Kabrick, J. M. Stand-density effects on aboveground carbon dynamics in secondary Pinus and Quercus forests of central USA. Sci. 69 (2), 213–227. https://doi.org/10.1093/forsci/fxac053 (2023).
Zhang, C. et al. Disturbance-induced reduction of biomass carbon sinks of china’s forests in recent years. Environ. Res. Lett. 10, 114021. https://doi.org/10.1088/1748-9326/10/11/114021 (2015).
Högbom, L. et al. Trilemma of Nordic-Baltic forestry – How to implement UN sustainable development goals. Sustainability 13 (10), 5643. https://doi.org/10.3390/su13105643 (2021).
Molina-Valero, J. A. et al. Mature forests hold maximum live biomass stocks. Ecol. Manag. 480, 118635. https://doi.org/10.1016/j.foreco.2020.118635 (2021).
Mikoláš, M. et al. Natural disturbance impacts on trade-offs and co-benefits of forest biodiversity and carbon. Proc. Biol. Sci. 288(1961), 20211631. https://doi.org/10.1098/rspb.2021.1631 (2021).
Wijas, B. J. et al. The role of Deadwood in the carbon cycle: implications for Models, forest Management, and future climates. Annu. Rev. Ecol. Evol. Syst. 55, 133–155. https://doi.org/10.1146/annurev-ecolsys-110421-102327 (2024).
Holzwarth, F., Kahl, A., Bauhus, J. & Wirth, C. Many ways to die – partitioning tree mortality dynamics in a near-natural mixed deciduous forest. J. Ecol. 101 (1), 220–230. https://doi.org/10.1111/1365-2745.12015 (2013).
Šēnhofa, S. et al. Availability and structure of coarse Woody debris in hemiboreal mature to old-growth Aspen stands and its implications for forest carbon pool. Forests 12, 901. https://doi.org/10.3390/f12070901 (2021).
Vasile, D. et al. Structure and Spatial distribution of dead wood in two temperate old-growth mixed European Beech forests. Not Bot. Horti Agrobot Cluj Napoca. 45 (2), 639–645. https://doi.org/10.15835/nbha45210829 (2017).
Lecina-Diaz, J., Senf, C., Grünig, M. & Seidl, R. Ecosystem services at risk from disturbance in europe’s forests. Glob Change Biol. 30 (3), e17242. https://doi.org/10.1111/gcb.17242 (2024).
Patacca, M. et al. Significant increase in natural disturbance impacts on European forests since 1950. Glob Change Biol. 29, 1359–1376. https://doi.org/10.1111/gcb.16531 (2023).
Ķēniņa, L., Elferts, D., Jaunslaviete, I., Bāders, E. & Jansons, Ā. Sustaining carbon storage: lessons from hemiboreal old-growth coniferous and deciduous forest stands. For. Sci. 69 (2), 158–166. https://doi.org/10.1093/forsci/fxac055 (2023).
Tíscar, P. A. & Lucas-Borja, M. E. Structure of old-growth and managed stands and growth of old trees in a mediterranean Pinus Nigra forest in Southern Spain. Forestry 89 (2), 201–207. https://doi.org/10.1093/forestry/cpw002 (2016).
Konôpka, B., Šebeň, V. & Pajtík, J. Species composition and carbon stock of tree cover at a postdisturbance area in Tatra National Park, Western Carpathians. Mt. Res. Dev. 39 (1), 71–80. https://doi.org/10.1659/MRD-JOURNAL-D-19-00008.1 (2019).
Bāders, E. et al. Norway Spruce survival rate in two forested Landscapes, 1975–2016. Forests 11 (7), 745. https://doi.org/10.3390/f11070745 (2020).
Krisans, O. et al. Presence of root rot reduces stability of Norway Spruce (Picea abies): results of static pulling tests in Latvia. Forests 11 (4), 416. https://doi.org/10.3390/f11040416 (2020).
Ahti, T., Hämet-Ahti, L. & Jalas, J. Vegetation zones and their sections in Northwestern Europe. Ann. Bot. Fenn. 5 (3), 169–211 (1968). http://www.jstor.org/stable/23724233
Lõhmus, A. & Kraut, A. Stand structure of hemiboreal old-growth forests: characteristic features, variation among site types, and a comparison with FSC-certified mature stands in Estonia. Ecol. Manag. 260 (1), 155–165. https://doi.org/10.1016/j.foreco.2010.04.018 (2010).
Rydin, H., Emanuelsson, U. & Kautsky, L. Biology and biogeographical zones of the Baltic region. In: Rydén L, Migula P, Andersson M (eds) Environmental science: understanding, protecting, and managing the environment in the Baltic Sea region. Baltic University Press, 68–91 (2003).
Bušs, K. Forest ecosystem classification in Latvia. Proc. Latvian Acad. Sci. 51, 204–218 (1997).
Aldea, J. et al. Species stratification and weather conditions drive tree growth in Scots pine and Norway Spruce mixed stands along Europe. Ecol. Manag. 481, 118697. https://doi.org/10.1016/j.foreco.2020.118697 (2021).
Ķēniņa, L., Elferts, D., Bāders, E. & Jansons, Ā. Carbon pools in a hemiboreal over-mature Norway Spruce stands. Forests 9, 435. https://doi.org/10.3390/f9070435 (2018).
Ķēniņa, L., Jaunslaviete, I., Liepa, L., Zute, D. & Jansons, Ā. Carbon pools in old-growth Scots pine stands in hemiboreal Latvia. Forests 10, 911. https://doi.org/10.3390/f10100911 (2019).
Sandström, F., Petersson, H., Kruys, N. & Ståhl, G. Biomass conversion factors (density and carbon concentration) by decay classes for dead wood of Pinus sylvestris, Picea abies and Betula spp. In boreal forests of Sweden. Ecol. Manag. 243 (1), 19–27. https://doi.org/10.1016/j.foreco.2007.01.081 (2007).
Liepa, I. Pieauguma mācība. Jelgava. (in Latvian) (1996).
Liepiņš, J., Lazdiņš, A. & Liepiņš, K. Equations for estimating above- And belowground biomass of Norway spruce, Scots pine, Birch spp. And European Aspen in Latvia. Scand. J. Res. 33 (1), 58–70. https://doi.org/10.1080/02827581.2017.1337923 (2017).
Liepiņš, J., Liepiņš, K. & Lazdiņš, A. Equations for estimating the above- and belowground biomass of grey alder (Alnus Incana (L.) Moench.) and common alder (Alnus glutinosa L.) in Latvia. Scand. J. Res. 36 (5), 389–400. https://doi.org/10.1080/02827581.2021.1937696 (2021).
Bārdule, A. et al. Variation in carbon content among the major tree species in hemiboreal forests in Latvia. Forests 12 (1292). https://doi.org/10.3390/f12091292 (2021).
Köster, K., Metslaid, M., Engelhart, J. & Köster, E. Dead wood basic density, and the concentration of carbon and nitrogen for main tree species in managed hemiboreal forests. Ecol. Manag. 354, 35–42. https://doi.org/10.1016/j.foreco.2015.06.039 (2015).
R Core Team. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from www.R-project.org. (2024).
Bates, D., Maechler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67 (1), 1–48. https://doi.org/10.18637/jss.v067.i01 (2015).
Kuznetsova, A., Brockhoff, P. B. & Christensen, R. H. B. LmerTest package: tests in linear mixed effects models. J. Stat. Softw. 82 (13), 1–26. https://doi.org/10.18637/jss.v082.i13 (2017).
Lenth, R. emmeans: estimated marginal means, aka least-squares means. R package version 1.5.5-1. (2021). https://CRAN.R-project.org/package=emmeans
Chai, Z. forestSAS: An R package for forest spatial structure analysis systems. R package version 0.1.0. (2016). https://github.com/Zongzheng/forestSAS
Zhao, Z. et al. Testing the significance of different tree Spatial distribution patterns based on the uniform angle index. Can. J. Res. 44 (11), 1419–0192. https://doi.org/10.1139/cjfr-2014-0192 (2014).
Szmyt, J. & Tarasiuk, S. Species–specific Spatial structure, species coexistence and mortality pattern in natural, uneven–aged Scots pine (Pinus sylvestris L.)–dominated forest. Eur. J. Res. 137, 1–16. https://doi.org/10.1007/s10342-017-1084-x (2018).
Lutter, R., Kõlli, R., Tullus, A. & Tullus, H. Ecosystem carbon stocks of Estonian premature and mature managed forests: effects of site conditions and overstorey tree species. Eur. J. Res. 138, 125–142. https://doi.org/10.1007/s10342-018-1158-4 (2018).
Kirschbaum, M. U. F. Is tree planting an effective strategy for climate change mitigation? Sci. Total Environ. 909, 168479. https://doi.org/10.1016/j.scitotenv.2023.168479 (2024).
Handegard, E., Gjerde, I., Bollandsås, O. M. & Storaunet, K. O. Identifying old Norway Spruce and Scots pine trees by morphological traits and site characteristics. Scand. J. Res. 36, 550–562. https://doi.org/10.1080/02827581.2021.1996628 (2021).
Šēnhofa, S. et al. Deadwood characteristics in mature and old-growth Birch stands and their implications for carbon storage. Forests 11 (5), 536. https://doi.org/10.3390/f11050536 (2020).
Mildrexler, D. J., Berner, L. T., Law, B. E., Birdsey, R. A. & Moomaw, W. R. Large trees dominate carbon storage in forests East of the cascade crest in the united States Pacific Northwest. Front. Glob Change. 3, 594274. https://doi.org/10.3389/ffgc.2020.594274 (2020).
Slik, J. W. F. et al. 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 (2013).
Bono, A. et al. The largest European forest carbon stocks are in the dinaric alps old-growth forests: comparison of direct measurements and standardised approaches. Carbon Balance Manage. 19, 15. https://doi.org/10.1186/s13021-024-00262-4 (2024).
Sims, A., Mändma, R., Laarmann, D. & Korjus, H. Assessment of tree mortality on the Estonian network of forest research plots. Forestry Stud. 60, 57–68. https://doi.org/10.2478/fsmu-2014-0005 (2014). https://intapi.sciendo.com/pdf/
Siipilehto, J. et al. Stand-level mortality models for nordic boreal forests. Silva Fenn. 54 (5), 10414. https://doi.org/10.14214/sf.10414 (2020).
Larson, A. J. et al. Spatial aspects of tree mortality strongly differ between young and old-growth forests. Ecology 96 (11), 2855–2861. https://doi.org/10.1890/15-0628.1 (2015).
Acknowledgements
The authors would like to thank to Annele Baltmane for her assistance with English language editing and grammar checking of the manuscript.
Funding
This study was supported by the European Regional Development Fund project “Development of a decision support tool integrating information from old-growth semi-natural forest for more comprehensive estimates of carbon balance” (No. 1.1.1.1/19/A/130) and the Latvia Council of Science national research programme project: “Forest4LV – Innovation in Forest Management and Value Chain for Latvia’s Growth: New Forest Services, Products and Technologies” (No. VPP-ZM-VRIIILA-2024/2–0002).
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Conceptualization: L.Ķ., Ā.J.; Methodology: Ā.J., L.Ķ., D.E., I.J.; Data collection: I.J., L.Ķ.; Formal analysis and investigation: L.Ķ., D.E., Ā.J.; Writing – original draft preparation: L.Ķ.; Writing – review and editing: L.Ķ., Ā.J., D.E.; Funding acquisition: Ā.J.; Supervision: Ā.J.
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Ķēniņa, L., Elferts, D., Jaunslaviete, I. et al. Disturbance event impact on aboveground carbon storage and vulnerability of large trees in old-growth coniferous forest stands in Latvia.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-37378-1
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DOI: https://doi.org/10.1038/s41598-026-37378-1
Keywords
- Stand density
- Carbon stock
- Deadwood
- Stand fragility
- Hemiboreal region
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